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Full text of "Memoirs of the California Academy of Sciences"

MEMOIRS 



OF THE 



^California Academy of Sciences 



Volume Vn 




Hydrographic History and Relict Fishes 

of the 
North-Central Great Basin 



BY 



CARL L. HUBBS, ROBERT RUSH MILLER, AND LAURA C. HUBBS 



SAN FRANCISCO 

PUBLISHED BY THE ACADEMY 

FEBRUARY 8, 1974 



MEMOIRS 



OF THE 



California Academy of Sciences 



Volume VII 




Hydrographic History and Relict Fishes 

of the 
North-Central Great Basin 



BY 



CARL L. HUBBS, ROBERT RUSH MILLER. AND LAURA C. HUBBS 



SAN FRANCISCO 

PUBLISHHD BY THE ACADEMY 

FEBRUARY 8, 1974 



Publication of this Memoir was made possible by a grant from the 

Belvedere Scientific Fund 



COMMITTEE ON PUBLICATION 

Al the time of ;icccptancc ol this nianiisciipl 

George E. Lindsay. Cluiiniuin 
Edvvai'd L. Kessel, Editor Leo G. Hertlein 



At the nortlu'tisi cud of this same raiii^t' of inoiiiitains [Riihy Raiii^c]. 
in the vcilley near the railroad town of Wells, are apparently bottomless 
fountains of water miles from any surface stream, ft is hut a few feet 
across the largest of them, the smaller con Id he crossed at a bound, and 
all are peopled with swarms of little fish, noiw of them over four inches 
in length. One hundred ami sevetity miles to the southwest are other 
wells in which can be found similar specimens of the finny tribe, but they 
e.xist nowhere else upon the continent. From what age, and condition of 
the past are they the relics? 

From Thompson and West's History of the State of Nevada. 1881. 
edited by Myron Angel, page 18. 



CONTENTS 



Introduction 1 

Field Surveys by Authors _ 2 

Quaternary Paleohydiography of Basins in the North-Central Great Basin 3 

Pluvial Lakes Regarded as More or Less Disjunct from the Lahontan System 10 

Pluvial Lake Gilbert _ _ 10 

Pluvial Lake Diamond _.. 15 

Pluvial Lake Diana 21 

Pluvial Lake Yahoo _ _ 21 

Pluvial Lake Newark _ 22 

Pluvial Lake Hubbs 26 

Pluvial Lake Clover _ 29 

Pluvial Lakes Regarded as Having Had Connections with Both Lahontan and 

Colorado Systems 32 

Pluvial Lake Railroad 32 

Pluvial Little Fish Lakes _., 35 

Pluvial Lake Lunar Crater 36 

Pluvial Lake Snyder 37 

Pluvial Lakes Containing the Relict Dace (between Lahontan and 

Bonneville Systems) . 38 

Pluvial Lake Franklin 39 

Pluvial Lake Gale 45 

Pluvial Lake Waring 46 

Pluvial Lake Steptoe 55 

Pluvial Upper Lake Steptoe 59 

Pluvial Lake Antelope 60 

Pluvial Lake Spring (Separated by Low Divides from Both Bonneville 

and Colorado Systems) 61 

Pluvial Lakes Regarded as Having Been Tributary to Colorado River 

System 64 

Pluvial Lake Carpenter 65 

Pluvial Lake Jake 66 

Pluvial Lake Pine C'Wah Wah") within Bonneville Svstem 68 



Fish Life of Basins in North-Centra! Great Basin _ 70 

General Treatment _ , 70 

Fish Fauna of Area: Its High Endemism and Extreme Depauperation 70 

Limited Sympatry _ . 76 

Areas of Basins Correlated with Richness and Diversity of Fish Faunas 77 

Characters and Systematic Status of Species 79 

Size 80 

Color. Texture. Form, and Sensory Structures 81 

Scale Structure 82 

Morphometry 82 

Sexual Dimorphism and Nuptial Characters 84 

Meristics 84 

Distribution and Habitat 95 

Structure and Status i^' Populations 96 

Speckled Dace. Rliiniclilliys osciiliis (Girard) 96 

Lahontan Speckled Dace, Rliiiiiclithys o. rohiisiiis (Rutter) 104 

Grass Valley Speckled Dace. R. o. rcUqiiiis Hubbs and Miller . 121 

Clover Valley Speckled Dace. R. o. oliiiopoyus Hubbs and Miller 129 

Independence Valley Speckled Dace. R. o. Icthoponis Hubbs and Miller .. 134 

Tui Chub. (Jihi hicolor (Girard) 142 

General Appraisal of Lahontan Subspecies 142 

Lahontan Creek Tui Chub. G. b. obcsa (Girard) 150 

Newark Valley Tui Chub. G. /'. Dcwaikcnsis Hubbs and Miller 156 

Fish Creek Springs Tui Chub. G. h. ciichila Hubbs and Miller 168 

Independence Valley Tui Chub. G. /'. isolata Hubbs and Miller .. 175 

Rcliiiii.s Hubbs and Miller 180 

Relict Dace. Rclicliis solilarius Hubbs and Miller . 196 

The Springfish. Genus Crciiichlhys Hubbs 227 

Railroad Valley Springfish. Crcnichthy.s iicvadac Hubbs 227 

Transfers of Great Basin Fishes (other than Game Species) into, between, and from 

the North-Central Basins 229 

Introductions of Exotic Fishes 235 

Survival and Conservation . 240 

Survival Status of the Endemic Fishes 241 

Hopeful Signs of Protection for Endangered Species of Desert Fishes 244 

Acknowledgments 244 

Literature Cited 246 

Index 254 



HYDROGRAPHIC HISTORY AND RELICT FISHES OF THE 
NORTH CENTRAL GREAT BASIN 

By 

Carl L. Hubbs, Robert Rush Miller, and Laura C. Hubbs 



INTRODUCTION 



As a belated sequel to our summary treatise 
(Hubbs and Miller, 1948b) that correlated in 
general terms the hydrographic history and the 
remnant fish life of the Great Basin and other 
arid parts of western North America, we now 
report, with more intensive documentation, on a 
cluster of more or less completely enclosed (en- 
dorheic ) basins in the center and very heartland 
of the Great Basin ( tables 1,2). This is a region 
wherein the geomorphic forces of the Basin and 
Range physiographic province have, during late 
Quaternary time, disrupted the topographic and 
drainage patterns to an almost unmatched degree. 
Here, bare remnants of fish fauna have survived 
in the extreme isolation that has resulted from the 
almost complete desiccation of an area that not 
more than a few thousand years ago was one- 
fifth covered by lakes fed by streams of ample 
flow. 

The area selected for present treatment in- 
cludes the contiguous drainage basins of 20 
pluvial lakes in central and eastern Nevada and 
of one lake, slightly disjunct, in western Utah 
(fig. 1 ). To the northwest, north, and east, the 
20 mostly separated basins in Nevada, are sur- 
rounded by the still largely integrated and much 
less fish-depleted Lahontan and Bonneville drain- 
age basins. To the north, these two main drain- 
age systems are narrowly conjoined, to separate 
the area being treated from the southernmost 
headwaters of the vast Columbia River system 
(the common divide between these drainages is 
reduced by a wedge of the Columbia River sys- 
tem to a straight-line distance of only 26 kilo- 



^ Conlribution from Scripps Institution of Oceanography, University 
of California, San Diego, and from the Museum of Zoology of The 
University of Michigan. 



meters). On the south side, the selected depres- 
sions in Nevada adjoin basins that in pluvial time 
were parts of the Colorado River system or were 
separated therefrom by low sills. Beyond the 
southwest boundary lie the intensely arid, fish- 
less "Area of Sterile Basins" and the basin of 
pluvial Lake Toiyabe (Hubbs and Miller. 1948b, 
pp. 45-51 and 44-45, respectively). Some en- 
dorheic basins have been somewhat arbitrarily 
excluded or included in our treatment, but we 
have maintained a set comprising, with a single 
exception, a contiguous area covering the north- 
central part of the Great Basin. This exception, 
that of pluvial Lake Pine (Wah Wah) in Utah, 
is wholly surrounded by the southwestern tribu- 
taries of the Bonneville system (p. 68). 

Together, the basins of these 21 selected 
pluvial lakes well represent the many endorheic 
valleys of the Great Basin as a whole. The 
selected area is indeed one of great ecological and 
fauna! stress. Next to the fishless "Area of Sterile 
Basins," it is characterized by the extreme dis- 
integration of drainage systems and by the almost 
maximal depauperation of the fish fauna, ac- 
companied by the extensive subspecific differenti- 
ation of the greatly isolated remnants of the 
fauna. 

In this report we deal with the fishes as well as 
the hydrography of all 21 basins, with the excep- 
tion that a detailed analysis of the fishes of the 
Lake Railroad system is reserved for later treat- 
ment. We have under preparation also an ac- 
count of the Lake Alvord system, astride the 
Oregon-Nevada line, and its two fish species. 
The cyprinodont fishes of the Death Valley sys- 
tem have been analyzed in detail (Miller. 1948. 
1961) and the paleohydrography of the region has 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



been treated (Miller. 1946: Morrison. 1965. fig. 
5). The fishes of the Lahontan system were 
provisionally reviewed l^y Snyder ( 1917). but the 
field exploration of the fishes of the Bonneville 
system that he conducted in 1915 never led to a 
thorough treatment of the fauna. Still earlier, 
Snyder ( 1908) had dealt with the fish fauna of 
the Oregon lakes. These and other early studies 
call for revision, and the limited faunas of a con- 
siderable number of endorheic basins south, west, 
and northwest of the north-central Great Basin 
area herein dealt with remain in need of treat- 
ment, though a brief summary of the hydrography 
and of the fishes has been presented ( Hubbs. 
1941a; Hubbs and Miller. 1948b). 

The studies that have Ixx^n published on the 
basins of the West and on their fish faunas, and 
the investigations herein reported, have been 
limited very largely to field reconnaissance, to 
the questioning of local residents, especially early 
settlers, to studies of available maps, and to the 
conventional systematic examination of the pre- 
served fish specimens. Much remains to be done 
along these lines, but other approaches need to 
be undertaken. Rewards may be anticipated from 
more intensive field studies of the basins, includ- 
ing precise levelling, from a thorough examina- 
tion of the aerial photographs now available, and 
through research on the subsurface lacustrine de- 
posits, to ascertain more .securely the past drain- 
age relations and the history of the ancient lakes. 
Similarly, more extensive and more reliable in- 
terpretation of the differentiating relict fishes of 
the sharply isolated remnant habitats may be ex- 
pected through such emergent areas of research 
as electrophoresis, karyotype analysis (beyond 
the start we have made since 1969), breeding and 
transference experiments, and through population 
and behavior studies. Any evidence — now totally 
lacking — on Cenozoic fossil fishes in the area 
herein covered would be enlightening ( p. 7.^ ). 

As we have indicated a number of times, the 
Great Basin is a vast arena wherein there has 
long been an active interplay between the pro- 
cesses of faunal establishment and extinctions. 



habitat disruption, and the isolation and dif- 
ferentiation of the remnant fish population. Docu- 
mentation of this interplay has been the key 
objective of this report. 

An abstract of this study has been published by 
us ( 1970). 

FIELD SURVEYS BY AUTHORS 

The authors" acquaintance with the portion of 
the Great Basin herein under treatment dates from 
1915 when the area was traversed by one of us 
(Hubbs. as student assistant to .lohn Otterbein 
Snyder), en route by car to and from a summer 
survey of the Bonneville system. The recognition 
of some of the waters in the basins as isolated 
would have led to the initial collection of some 
of the fishes herein treated, had it not been for an 
emergency that forced return without delay. 

Together. o\- separately, the authors carried on 
field work in the area and in related parts of the 
Great Basin in 1*^)26. followed by major trips in 
1934. 1938. and 1942. with briefer supplemen- 
tary studies in various years from 1950 to the 
present. 

Our field procedures included relentless ques- 
tioning of informed local residents, including, by 
good fortune, a number of original settlers and 
their children, who aided greatly in orienting us 
at a time when mapping was crude and limited, in 
providing history on the rare flash floods of the 
desert region, in telling us which of the isolated 
waters and basins did or did not contain fish, and 
in clarifying which species were native and which 
were introduced. With such help we crossed and 
recrosscd nearly all of the basins under discussion 
and various adjoining basins. We have felt rather 
confident that we have sampled nearly all of the 
springs or groups of springs containing native fish. 
Shoreline features of ancient lakes were diligently 
sought, and their altitudes were checked against 
possible outlet passes by use of Paulin precision 
altimeters. Other pertinent geographical and 
geological features were observed and recorded. 
.An effort in 143X to obtain data on the chemical 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



composition of the often highly mineraUzed waters 
of isolated springs unfortunately proved abortive 
(p. 95 ) . Large samples of the native fishes were 
collected and carefully preserved, with extensive 
geographical and ecological notes. Through the 
years these specimens have been studied exten- 



sively, in comparison with material from various 
other Western waters. Many colleagues ( pp. 244- 
246) have aided us with geological, ecological, 
and ichthyological data and advice. 

The preparation of this report, with help from 
many, has been underway since the end of 1964. 



QUATERNARY PALEOHYDROGRAPHY OF BASINS 
IN THE NORTH-CENTRAL GREAT BASIN 



The basins of the 2 ! selected pluvial lakes 
typify the many more or less disrupted fault-block 
depressions that feature the topography of the 
Basin and Range physiographic province, be- 
tween the east-facing escarpment of the Sierra 
Nevada and the west-facing front of the Wasatch 
Range (Fenneman, 1931; Nolan, 1943; Black- 
welder, 1948; Hubbs and Miller. 1948b; King, 
1958; Snyder, Hardman. and Zdenek. 1964; Feth, 
1 964; Morrison, 1 965 ) . Most of these basins and 
the beds of their contained lakes are elongated in 
a more or less exactly north-south direction. They 
are bounded on one or both sides by great fault 
scarps, along which typically issue the valley- 
bottom or valley-edge .springs in which, often al- 
most unbelievably, a remnant of the fish life of 
the basin has somehow persisted. In contrast, the 
canyon and mountain-side springs of the basins, 
though seemingly permanent and hospitable, 
.seldom contain native fish, presumably because 
occasional great flash floods, induced by the rare 
torrential precipitation of the desert region, sweep 
everything before them, then subside too rapidly 
to allow repopulation by return dispersal from 
the playa below. 

Although many of the basins of the area have 
probably existed since at least late Miocene time, 
long enough to render plausible the retention of 
a few localized, trenchantly distinct, relict fishes 
(p. 70), most of the perceptible topographic 
and ichthyological evidence seems to pertain to 
post-Sangamon time, and much of it to the late 



Wisconsin. Most of the earlier physiographic 
evidence of ancient lakes and streams has been 
eroded away, and probably most of the fishes 
that earlier occupied the area have failed to sur- 
vive. In our previous discussion (Hubbs and 
Miller, 1948b). we formalized the late events 
that are still signalized by ancient beachlines and 
other obvious physiographic evidence by referring 
to them, and to the lakes and streams that they 
represent, as Pluvial (with initial capital), but 
the suggestion has not been followed and we now 
abandon the practice. 

We are also altering our former terminology in 
the naming of the pluvial lakes. Previously, we 
followed for most of these lakes the format of 
putting the name 'Lake" first, following the pre- 
cedent of Lake Bonneville, Lake Lahontan. Lake 
Manly, Lake Algonquin, etc., but reversed the 
word order for certain basins when the combina- 
tion seemed somewhat awkward or possibly con- 
fusing. Thus we used, among the lakes herein 
considered, Antelope Lake. Pine Lake. Railroad 
Lake, and Spring Lake. We now think it better 
to adopt the names Lake Antelope. Lake Pine, 
Lake Railroad, and Lake Spring, and the like, 
feeling that the consistent use of capital letters 
will preclude the double meaning. For ancient 
Upper Steptoe Lake we alter the word order to 
Upper Lake Steptoe. For the relatively insignifi- 
cant ancient version of Little Fish Lakes we do 
not alter the word order. 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



The highest level and the discharges of some 
of the ancient lakes are indisputably demonstrated 
by beachiines and other evidence, but other con- 
nections remain more or less hypothetical (as is 
indicated in the detailed accounts that follow ) . In 
such cases, considerable weight has been given to 
the fish evidence, for the natural occurrence of the 
same or of very closely allied fish types is very 
strong evidence of a not very remote past water 
connection. Also, when clear evidence of beach- 
lines is seen by field reconnaissance, or by the 
examination of the excellent topographic maps 
that have recently become available, to lie not 
far below the level of the assumed outlet pass, we 
venture the assumption that during occasional 
periods of unusually heavy precipitation the lake 
rose high enough to discharge. Anyone who has 
observed ordinarily dry playas of ancient lakes 
fill suddenly to a considerable depth by bursts of 
torrential precipitation, or to retain water for a 
considerable time in wet years, as in 1965, is not 
likely to doubt that during especially moist years 
such pluvial lakes overflowed at least briefly. 
Furthermore, the shoreline features of relatively 
early outlet levels and of temporary later ones 
may have been obliterated by time. We have 
relied on such reasoning to counterbalance the in- 
complete evidence for lake levels not quite high 
enough to insure discharge, particularly for lakes 
Gilbert (p. 11), Newark (p. 23), Carpenter (p. 
65), and Jake (p. 67). For Lake Franklin (p. 
42) and probably for Lake Clover (p. 30), at 



latest stages, we find indications of a subterranean 
discharge through alluvium. 

In interpreting the hydrographic relations of 
the pluvial waters under consideration ( tables 1 , 
2) and in their delineation (figs. 1-3, 7, 8, 11- 
14), we have relied largely on our earlier review 
( Hubbs and Miller, 1948b) and on the annotated 
map by Snyder ct al. ( 1 964), which was essentially 
copied by Feth (1961, 1964) and Morrison 
( 1965. fig. 1 ), and, of course, on various earlier 
workers cited in these treatises. We have reviewed 
the field notes of our several reconnaissances, 
during most of which we used a precision Paulin 
altimeter. Our field trip in 1965 clarified the 
shoreline features of pluvial lakes Clover, Frank- 
lin, and Hubbs. 

During recent years, even after the appearance 
of the new maps of Great Basin lakes just cited, 
a number of geologists have briefly treated these 
lakes, largely on the basis of the account and 
small-scale map by Flint (1957, pp. 226-233, 
fig. 13.2). Among these authors we cite Strahler 
(1963. pp. 357-358, fig. 31.25); Shelton (1966, 
pp. 352-363, figs. 331-343); Engel ( 1969, pp. 
476-477, fig. 10); Longwell, Flint, and Sanders 
( 1 969. pp. 280-28 1 . fig. 1 2.28 ); and Thornbury 
(1969, pp. 4(W-407, figs. 16.13, 16.14). Flint 
(1971. pp. 442-451, figs. 17.1-17.5) has further 
treated the lakes. A close approach to our present 
interpretations was issued by Morrison (1965, 
pp. 265-285, fig. 1 ), with a review of the deposi- 
tional data, which were further treated by Mor- 
rison and Wrieht ( 1967). 



Figure I. Guide map to hydrographic features discussed in this report, and in adjacent areas; adapted from the 
map of Pleistocene lakes in the Great Basin, by Snyder, Hardman, and Zdenek ( 1964). 

Indicated are areas, in clockwise sequence. D. C, A, B, as covered by the four pairs of detailed maps, respectively 
pluvial (figs. 2, 7, 11. 13) and modern (figs. .3. 8. 12, 14). Shown also are all recognized pluvial lakes in the area 
(named in sloping type, with new designations underlined); also many streams (major flows, and others helpful in 
showing drainage relations), and major towns. Stars designate the 21 pluvial lakes particularh' treated in this report. 
The black center of Lake Steptoc represents the pluvial size that now seems to us most plausible. Otherwise, lake 
areas are essentially as mapped by Sn\der ci al.. generally in close agreement with us, except that we recognize, for 
a few lakes, somewhat hypothetical, higher maximum outlet levels (these outlets are depicted on this map, although 
the lake levels as charted would have been below spillage). Question marks denote two Pleistocene lakes that were 
recognized by Snyder ci al. ('Hot Creek Lake" and one in Egan Valley on the western side of the Steptoe basin). 
but not accepted by us, at least as of pluvial age. RO and Rl represent collections of Rhiiuchthys iiscuhis rohiistiis. 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table I. Busic ilntn <>n the 21 jtluv'ui! hikes in lunlh-cenlrcil Great Bashi tieateil in this report. 



Pluvial 

lake name 
(alphabetical )- 


Vallev 
(for lake) 


Area on 
fig. 1 


Map : 
fig. 


County 
(for lake)' 


Discharged?' 


Tributary to' 


Antelope (29, 8=) 


Antelope 


A. B 


11, 13 


White Pine 


7 


L, Waring (?) 


Carpenter (66, 50) 


Lake (Duck) 


S of B 


13 fN. end) 


Lincoln, White 
Pine 


Yes 


Carpenter R. & 
Colorado R. 


Clover (26, 15) 


Clover, 

Independence 


A, C 


7, 11 


Flko 


Early" 


Humboldl R. & 
1. Lahontan 


Diamond (4, 68) 


Diamond & Kobeh 


C, D 


2. 7 


Fureka, Flko 


Yes 

(early'') 


Humboldt R. & 
L. Lahontan 


Diana (— , 60) 


Monitor 


D 


2 


Nye 


'ies 


Monitor R. & 
L. Diamond 


Franklin (24, 38) 


Ruby & Butte (N) 


A, C 


7, 11 


Flko. White Pine 


■>» 


L. Clover (?) 


Gale (25, 107) 


Butte (Si 


A. B, C 


7, 11. 13 


White Pine, Flko 


Yes 


L. Franklin 


Gilbert (5, 70) 


Grass 


D 


T 


Lander, Fureka 


'les 

(early'') 


Humboldt R. & 
L. Lahontan 


Hiibbs (?1, 52) 


Long 


C 


7 


White Pine 


Early?'? 


L. Newark (??) 


Jake (,^2, 45) 


Jakes 


B, C 


7, 13 


White Pine 


Yes 


White R. & 
Colorado R. 


little Fish (— , 51) 


little Fish V. 


O 


T 


Nye 


Yes 


Russell R. & 
L. Railroad 


lunar Crater 
(— . 94) 


Sand Spring 


S of D 


— 


Nye 


? 


L. Railroad (?) 


Newark (3. 64) 


Newark 


C, D 


2, 7 


White Pine, 
Fureka 


Yes 

(early?) 


Humboldt R. & 
L, Lahontan 


Pine (Wah Wah ) 
(If, 88) 


Pine 


SF of B 


— 


Millard, Beaver 
(Utah) 


No 


Surroimded by 
Bonneville 
basin 


Railroad (60, 80) 


Railroad ct ul. 


D + SE of 
D 


T 


Nye 


Very early 


White R. & 
Colorado R. 


Snyder (— , 106) 


(See text) 


D 


-» 


Nye 


Probably 


Lunar Crater L. 
& L. Railroad 
(?) 


Spring (30, 98) 


Spring 


A, B 


11, 13 


White Pine 


77 


L. Bonneville 
(?) or L. Car- 
penter . . . 
Carpenter R. 
& Colorado 
R. (?) 


Steploc (27, 99) 


Steptoe ( north ) 


A. B. C 


7, 11, 13 


While Pine. Flko 


Yes 


L. Waring 


Upper Steptoe 
(— , 100) 


Steptoe (south) 


B 


13 


White Pine 


^'es 


L. Steptoe 


Wariny (28, 38) 


Goshute 


A. B 


11. 13 


Flko 


No 


Nearest L, 
Bonneville 


Yahoo (— , 81 ) 


■\ahoo 


C, D 


2, 7 


Fureka 


Yes? 


Monitor R. & 
L, Diamond? 



' Supplementary to table 2. 

-The lakes constituting the block treated are marked with an asterisk on the guide map (tip. I), lo designate the area covered. 

' All m central and eastern Nevada, except for Pine Lake, in Utah. 

'At maximum lake level {in part somewhat hypothetical). 

f^'The numbers in parentheses are, respectively, those used on the maps of Hubbs and Miller tI'^4Sb) and of Snyder ef at. (1964). 

" Discharge postulated as subterranean during late pluvial time. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



Table 2. Drainai;e relations, native fish species, depth, and areas of the 21 piiivicd lakes treated.^ 



Pluvial lakes" and drainage relations 



Native 

fish 
species' 



Est. max. 
depth, m. 



Area of lake 



Lakes regarded as more or less disjunct from 
the Lahontan system 



Gilbert (5. 70)" 

Diamond | 

complex 



Newark 
complex ( ? ) 



Diamond (4. 68) 

Diana (— . 60) 
Yahoo (— . 81) 

Newark (3. 64) 



Hubbs (31, 52) 
Clover (26. 15) 
Lakes regarded as having had early connec- 
tions with both the Lahontan and Colo- 
rado systems 



Railroad 
complex 



Railroad (60. 90A. B) 



G. R 



G. R 



Little Fish (— . 51) 
Lunar Crater ( — . 94) 
Snyder (— . 106) 
Lakes containing the relict dace, in enclosed 
basins between the Lahontan and Bonne- 
ville systems 

Franklin f Franklin (24. 92) 

complex \ Gale (25. 10) 

r Waring (2S. 38) 
Waring | Steptoe (27. 99) 

complex 1 Upper Steptoe ( — . 100) — 

I Antelope (29. 8) 
Lake separated by low divides from the 
Bonneville and Colorado systems 
Spring (30. 98) 
Lakes regarded as having been tributary to 
the Colorado River system 

Carpenter (66, 50) 

Jake (32. 45) 

Lake enclosed within southwest corner of 
Bonneville system 

Pine (Wah Wah) (\c. 88) 



r 56 

"1 79 


478 - 
565 - 


- 1.528 = 31%" 

- 1,528 = 37%''' 


r 45 

1 76 
shallow 
shallow 


792- 
1.002- 

12- 
7 - 


- 8,097^ = 10%" 

- 8,097' = 12%" 

- 1,339 = 1% 
-44= 16% 


r 69 

1 115 

63 


792- 

1.146- 

490- 


- 3.587 = 22%" 

- 3.587 = 32%" 

- 1.677 = 29% 


42 


930 - 


- 2.624 = 35% 



C G 


104 / 


1,394 - 


- 10.874^= 13% 






1.394- 


- 12.343"= 11% 


G 


shallow 


11 - 


- 1.131 = 1% 


— 


shallow 


1 1 - 


- 1.321 = 1%, 


— 


shallow 


24 - 


- 148 = 167^ 



S 


53 


1.314- 


-5,271" = 25% 


s 


35 


474- 


- 1,933 = 257r 


S 


66 


1.314- 


- 9,411" = 14% 


s 


<30 


282 - 


- 5,462'-" = 5% 


— 


shallow 


13 - 


- 816 = 2% 


— 


23 


125 - 


- 870 = 14% 



95 



978 -:- 4,337 = 23% 



<26 


247 ^ 1,257 = 20% 


33 


186 -^ 1.114 = 17%° 


46 


249- I.I 14 = 22%" 


60 


102^ 1,912 = 5% 




GRAND TOTALS 



Total lake areas' 



Area' of drainage basin 



Area' of entire basin 



1.021 



1,282 
1.636 



8,097 = 107c" 
8.097= 137<," 



5.264 : 
5,264 : 



: 24%" 

:31%.« 



1.440 ^ 12.343 = I27o 



1.788 -^ 5.271 = 34%, 



1.734-:- 10.281 = 17%- 



433 
496 



2.371 = 18%>° 
2.371 =2170" 



f 9,976 -f- 54,028 = 18.57o" 
\ 10,690 ^ 54,028 = 19.87,," 



1 Supplementary to table !. 

-The 21 pluvial lakes treated in this report are marked with an asterisk on the guide map (fig. I), to designate the area covered; all may not 
have been strictly contemporaneous. 

3 C, Crenichthys nevadae; G, GiUi bicolor subspecies; R. Rhinichthys oscnltis subspecies; S. Reluttis solitaihis. 

' All areas given in square kilometers. 

^'The numbers in parentheses after the lake names are those used, respectively, by Hubbs and Miller (1948b) and by Snyder et al. (1964). 

*> For lakes Gilbert, Diamond, Newark, and Jake we give values for two estimates of the maximum level: first, one hardJy subject to doubt; 
second, one based on the assumption of a discharge level, perhaps not attained during late pluvial time. 

'Drainage area including basins of lakes Diana (evidence clear) and Yahoo (evidence unclear). 

* Drainage area of Lake Railroad and the clearly tributary Little Fish Lakes, but not that of lakes Lunar Crater and Synder. 

^ Area including the dubiously tributary drainage basins of lakes Lunar Crater and Snyder. 

I" Including the certainly tributary drainage basin of Lake Gale. 

'1 Including the certainly tributary drainage basins of Lake Steptoe and Upper Lake Steptoe. 

'^ Including the certainly tributary drainage basin of Upper Lake Steptoe. 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



or inestimable help have been the magnifi- 
eently detailed and accurate topographic maps 
published in recent years by the United States 
Geological Survey (how these would have helped 
us in our earlier studies!). The entire area has 
been covered by 1 5 National Topographic Maps 
1 :25(),0()()-scale located as follows, from north 
to south, in each of three vertical rows (from 
west to east), with dates of survey or of limited 
revision: (1) McDermitt 1959, Winnemucca 
195S, Millctt 1959, Tonopah 1959, and Gold- 
field 1958; (2) Wells 1958, Elko 1958, Ely 
1959, Lund 1960, and Caliente 1959: (3) 
Brigham City 1958, Tooele 1958, Delta 1962, 
Richfield I 958, and Cedar City 1958. 

The few then available earlier tiipographic 
maps, some on the 30-minute scale, were useful 
during the field work and still have some interpre- 
tive value. For historical purposes, as in tracing 
early explorations and original conditions, con- 
siderable use was made of old maps, including 
those in the reports of the Commissioner of the 
General Land Office for 1 862 and, more particu- 
larly, for 1866 (see p. 233) — covering the early 
exploration of Nevada. Useful also, in the same 
connection, has been a copy of a map published 
during early years of mining entitled: 

Map of the States of California and Nevada, 
carefully compiled from the Latest Authentic 
.Sources, By Chas. Drayton Ciihbes, C. E. 
Comprising Information obtained from the 
U. S. Coast and Land Surveys, State Cieolog- 
ical Surveys, by Prof. J. D. Whitney, Railroad 
Surveys, and the Results of Explorations Made 
by Brevet Lieut. Col. R. S. Williamson, U.S.A., 
Henry tie CJroot, C. D. Gihbes, and Others. 
Published by Warren Holt. I,S7.V No. 607 
Cla\ Street. San Francisco, Caj. Scale 18 
miles to OIK inch. 

This map was indicated as having been drafted 
and lithographed by S. B. Linton, formerly of the 
U. S. Coast Survey. A much reduced copy was 
reproduced in the book by Wheeler ( 1 97 I , op- 
pt)sitc page 108). We have a photocopy of the 
original provided by the Bancroft Library. We 
have also checked the considerably detailed map 



of Nevada on pp. 830-83 1 of the Rand, McNally 
& Co.'s Indexed Atlas of the World. 1881 (in the 
senior author's library ) . 

The paired modern and pluvial maps ( repro- 
duced as figs. 2, 3, 7, 8, 1 1-14) were sketched 
on and traced from these 1:250,000 maps, uti- 
lizing all evidence available to us. Where, as was 
true for a large share of the area, 15-minute topo- 
graphic quadrangles had become available, the 
pertinent modern and ancient features were 
marked thereon and transferred to the 1 :250,000 
scale by a preci.se reducing apparatus. These 1 5- 
minute quadrangles were available for the entire 
area covered by 42 maps between 39 00' and 
40 45' N. lat. and between 115" 30' and 117° 
00' W. long., and for parts of the region east and 
southeast of those limits, for which the following 
sheets have been available ( listed from north to 
south in 7 vertical rows, from west to east, the 
first row below the easternmost column of the 
main block): ( 1 ) Duckwater 1964, Blue Eagle 
Springs 1964, and Troy Canyon 1964; (2) 
Lamoille 1962, illipah 1 95 1 , Treasure Hill 1949, 
Currant Mountain 1957, Currant 1964, and 
Forest Home 1964; ( 3 ) Riepetown 1959, Preston 
Reservoir 1959; (4) Spruce Mountain 1953, 
McGill 1958, and Ely 1958: (5) Spruce Moun- 
tain 4 1959, Schell Peaks 1959, Connors Pass 
1959; (6) Sacramento Pass 1958, and Wheeler 
Peak 1948; (7) Garrison 1959 (see Index to 
Topographic Maps of Nevada ). Within the Riepe- 
town, Preston Reservoir, McGill, and Ely 15- 
minute quadrangles some 7.5-minute quadrangles 
have been available and have been used to check 
some features, particularly where in this area 
mining operations have obfu.scated the topogra- 
phy. Here, an effort was made to portray the 
original condition. Use was also made, especially 
in our earlier work, of various other maps, in- 
cluding United Stales Geological Survey state 
maps, the United States General Land Office map 
of the State of Nevada ( 1930), United States 
Forest Service maps, and privately published 
county maps. Particularly helpful in the early 
field work was a map of White Pine County (Ed. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



Millard & Son. Ely. 1930). Recently there has 
become available a vastly improved and very de- 
tailed United States Geological Survey map of 
the State of Nevada, 1:500,000 (compiled 1962, 
edition of 1965). 

The topographic maps were especially valuable 
in tracing the divides between the various drain- 
age basins, which were shown rather crudely on 
our 1948 map, drawn when very few such maps 
were available, and these fell far short of the high 
modern standards of cartography set by the new 
topographic maps. In this respect, even the map 
by Snyder et al. ( 1964) is somewhat generalized. 
The boundaries of the many drainage basins of 
the state have been delineated in fine detail on the 
map of State of Nevada Water Resources and 
Inter-basin Flows (Rush et al., 1971), but on 
the basis of the modern rather than the pluvial 
drainage system. 

The areas of the drainage basins were mea- 
sured by planetary planimeter. The area in 
square kilometers (sq. km.) was determined after 
multiple measurements had been made of the 
areas in question and of 100 sq. km. on the 1: 
250,000 maps. 

The pluvial lake margins were drawn largely 
by following a contour, or by interpolation be- 
tween contours, on the most detailed topographic 
maps available, after points were fixed from field 
reconnaissance and map study and by comparison 
with the lake outlines as depicted by Snyder el al. 
( 1 964 ) . In general, our conclusions agree closely 
with theirs. In utilizing contours, we have relied 
on the large mass of evidence that the ancient 
shoreline features have very seldom been con- 
siderably distorted during the geologically short 
time that has elapsed — about 10,000 to 30,000 
years — since the still readily recognizable shore- 
line features were carved. In mapping the west- 
ern shoreline of Lake Bonneville (figs. 11, 13), 
reliance was placed on new data on the isostatic 
deformation of the basin since the evaporation 
of that great inland sea ( Crittenden. 1 963 ) . 

Each of the 21 pluvial basins under treatment 
embraced a pluvial lake, ranging in area from 



about 7 to 1,394 sq. km., of which 10 (Gilbert, 
Diamond, Newark, Hubbs, Clover, Railroad, 
Franklin, Gale, Waring, Spring ) covered at least 
approximately 500 sq. km. (table 2). Our two 
estimates for the total area of the 21 lakes, at 
maximum level, are 10,690 or 9,976 sq. km. 
( using two shoreline estimates for the area of 
lakes Gilbert, Diamond, Newark, and Jake ) : 1 9.8 
or 18.5 percentage of the total estimated area 
(54,028 sq. km. ) of the combined drainage area. 
Except for 6 very small and mostly stream-course 
lakes, listed as shallow, all of the 2 1 lakes are 
estimated to have reached depths between 23 and 
115 m. (table 2). The total storage of surface 
water must have been immense — surely more than 
100-fold greater than at present. The same can 
safely be said also of the differential in the propor- 
tion of the surface area that was covered by water. 
The two factors thought by some to have been 
adequate for the accumulation of surface waters 
during the Pleistocene, namely cooler tempera- 
tures and reduced evaporation, seem impotent, 
in themselves, to have produced the observed 
Pleistocene effect. For example. Cole ( 1968, p. 
427 ) stated that ". . . . the existence of Pluvial 
lakes that are now ephemeral playas .... de- 
pended on lower mean temperatures, reduced 
evaporation, and the resultant increased runoff." 
With others, we feel that markedly increased 
precipitation must also have been involved — 
presumably in addition to and in correlation with 
these other factors. Snyder and Langhein ( 1962. 
p. 2394) thought it highly implausible that either 
an increase in precipitation or a decrease in evapo- 
ration could alone have caused the accumulation 
of Lake Spring; they stated: "the combination 
of an increase in precipitation from the present 12 
to 20 inches [30-51 cm.] and a reduction in 
evaporation from 44 to 31 inches [112-79 cm.], 
that would restore Pleistocene Spring Lake, repre- 
sents a most probable combination on a statistical 
basis." These estimates probably apply reason- 
ably well to the north-central Great Basin as a 
whole. 

The climatic basis for the accumulation of vast 



10 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



bodies of watci in the Great Basin lias been ex- 
pressed, eonsimant witii our thinking, by a lead- 
ing student of the Quaternary geology of the 
region ( Morrison. 196?. p. 267 ) as follows: 

.... the Pleistocene clim.ite tlucUiatcd widely, 
with respect to both teiiipcrature and precip- 
itation. I he cooler and wetter times . . . called 
pliiviaK .... were .... 4.4 to 8..^° C cooler 
than now. and also appreciably less arid, as 
attested, tor example, bv markedly increased 
supply of coarse alluvium I n>m the mountains 
and b\ increased mass-wasting. Runoff in- 
creased because of lower temperature and 
greater precipUation. tipping the balance be- 
tween inflow and evaporatiem ni the basins in 
favor ot inflow, so that permanent lakes de- 
veloped in all the terminal basins. These "lake 
cycles" or "laerrstral intervals" were synchro- 
nous with intervals ol glaeiatron rn the higher 
mountains. 

PLUVIAL LAKES REGARDED AS MORE 

OR LESS DISJUNC T FROM THE 

LAHONTAN SYSTEM 

Seven of the 21 pluvial lakes under present 
study were in basins that are physiographieaily 
most elosely tied to the Humboldt River division 
of the Lake Lahontan systetii. and most, perhaps 
all of these, were, we think, at some pluvial 
period actually within that drainage system. The 
four main lakes involved were Gilbert, Diamond, 
Newark, and Clover. Two very minor lakes, 
Diana certainly and 'S'ahoo c|uestionably, we as- 
sociate with the Lake Diamond drainage system. 
The seventh lake of the series. Lake Hubbs, did 
not rise high enough, according to any definite 
evidence, to spill into Lake Newark. 

Pluvial Laki-: Gilbert 

Drainage basin on either side, near the middle, 
of the north-south line forming the boundary be- 
tween Lander and Eureka counties ( with the 
greater area in Lander County), in central 
Nevada (figs. 1.2). 

This rather large bodv of water was named 



pluvial Lake Gilbert by us (Hubbs and Miller, 
194Kb, pp. .^3-36, 148, 157). "in memory of 
Grove Karl Gilbert's classical contributions to 
Great Basin hydrography." This name has been 
accepted by Snyder el iil. ( 1964) and by Feth 
(1964). Lake Gilbert covered the large flat of 
Grass Valley and the lower levels of the adjoining 
bajadas. The middle of the valley toward the 
north end is occupied by a level, bare, sandy dry 
lake measuring 23 ■ ca. 3 km. on the Millett and 
Winnemucca 1:250,000 maps. The playa is 
rimmed, especially on the east side and around 
the north end. by sand dunes, some of consider- 
able size. The south end of the valley, which is 
bordered on each side by high mountains, Toiyabe 
Range to the west and Simpson Park Mountains 
to the cast, is less arid than the main part of the 
basin. 

The drainage basin, that of Grass Valley ( using 
this name, as is customary, to cover the whole 
basin, not merely the largely enclosed, more 
mcadcnvy southern arm), is bounded as follows: 
at the north tip. in the immediate vicinity of the 
outlet. Cortez (Tenabo) Canyon, by the drain- 
age basin of Crescent Valley, site of Lake Cres- 
cent (of uncertain age and status, for the 
valley maintains a flood-time outflow into Hum- 
boldt River — see p. 107); on the east side 
by the watersheds of Pine C tcek. of the 
Humboldt River system, and b\ the drain- 
age basins of Kobeh and Monitor valleys of the 
pluvial Lake Diamond system; on the south by 
the drainage basin of Big Smoky Valley, the site 
of pluvial Lake Toiyabe; on the west for a short 
distance by tributaries of Reese River, of the 
Humboldt River system, and for a greater distance 
by the watershed of C arico Lake Valley, which, 
like Crescent Valley, now is mapped as having 
a through drainage, but has been held to contain 
the bed of pluvial Lake Carico. 

The drainage basin trends north-northeast to 
south-southwest. It is oblong, with a triangular 
tip at the northern, outlet end. 

The basin measures 69 km. in greatest length 
and 32 km. in maximum width (at the north 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



11 



end of the oblong area, most of which is about 
25 km. wide). The lake was elongate, with a 
triangular expansion on the west side. We esti- 
mate its maximal length as 54 km. and its maximal 
width, at the submedian expansion, as 1 8 km. 

We compute the area of pluvial Lake Gilbert 
at its assumed outlet level as 565 sq. km., ap- 
proximating 37 percent of the area (1,528 sq. 
km. ) of its watershed. The computations for the 
more completely authenticated, late Pleistocene 
lake level are: area 478 sq. km., 3 I percent of the 
area of the watershed. At this lesser size ( indi- 
cated by a dashed line on fig. 2) the primary 
dimensions would be 17 -45 km. 

Inasmuch as Lake Gilbert lay in the latitudinal 
belt of deep pluvial lakes, and as its rim rose to 
heights as great as 920 m., the great extent of 
this ancient lake and its probable attainment of an 
outlet are consistent with expectation. 

The physiography and hydrography of this 
basin have been treated by Everett and Rush 
( 1966) who mapped the lake below a discharge 
level. 

Shoreline and Discharge. 

Our reconnaissance of August 9, 1938. led 
us to believe that Lake Gilbert discharged north- 
ward into Crescent Valley, by way of a steep 
canyon, named Cortez or Tenabo. Above the 
dunes margining the dry lake, the bajada is 
marked on each side by terraces that cut trans- 
versely across draws and alluvial cones. As seen 
from Walti Hot Springs, on one of the lower ter- 
races just east of the playa, a terrace on the west 
side seemed to line up with Cortez Pass, below 
which Cortez Canyon drops steeply into Crescent 
Valley. Altimeter readings indicated a succe.ssion 
of apparent beachlincs from about 100 feet (ca. 
30 m.) below to 100 feet higher than Walti Hot 
Springs. The uppermost terraces were rather 
faint, but seemed definite. A reading taken within 
an hour at the summit of Cortez Pass was only 10 
feet (3 m. ) higher. Near Cortez a broad saucer- 
like flat area, circling the north rim of the valley, 
is covered by seemingly lacustrine clay almost to 



the summit of the pass. It is assumed that at some 
pluvial time(s) Lake Gilbert spilled, but it does 
not appear to have long retained discharge during 
late pluvial time. The occurrence in the spring 
at Grass Valley Ranch of a very strongly modified 
form of the Rhinichthys osculiis group ( see be- 
low) is consistent with this view. 

The depth of Lake Gilbert was estimated by 
Snyder cl al. ( 1964) as 250 feet (76 m.). We 
attain estimates of 258 feet (79 m. ) for the as- 
sumed outlet level and 183 feet (56 m.) for the 
more conservative and less questionable late 
Pleistocene level on the following bases. Several 
altitudes are given on the Cortez and Walti Hot 
Springs 15-minute quadrangles for the playa level 
as 5,617 feet ( 1.712 m.), and one is marked as 
"VABM"" (Verified Altitude Bench Mark). The 
outlet level is taken as approximately 5,875 feet 
(1,791 m. ) because the paired 5,900-foot ( 1 ,798- 
m. ) contours run through the gentle pass and 
altitudes of 5,856 and 5,865 feet (1,785 and 
1,788 m.) are given just above the 5,850-foot 
( 1,783-m.) contours that cross the divide at the 
south and north sides. The less questionable and 
presumably later level is estimated, and mapped, 
as 5,800 feet (1,768 m.), because apparently 
clear evidence of lake terraces were noted as being 
100 feet above Walti Hot Springs, which are 
mapped as issuing on the 5,680-foot ( 1,731-m.) 
contour (sec cover photograph. Everett and 
Rush, 1966). 

Both estimated lake borders (fig. 2) are 
mapped by utilizing the following 15-minute 
quadrangles: Cortez 1938, Hall Creek 1956, 
Walti Hot Springs 1956, Mount Callaghan 1956, 
and Ackerman Canyon 1956. These and the 
other available, pertinent quadrangles were used 
in delineating the divides that surround the basin. 

The more profuse waters of possibly early 
pluvial times are indicated not only by high 
shoreline terraces and presumed discharge, but 
also by the strength and height of the stream ter- 
races along the draws fanning into the southern 
end of Grass Valley. These terraces seem incom- 
patible with the present stream flows, as do the 



12 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




r -i 


_. /---:---\ r- 


/ ^, 




/ -, 


— '1 <i \ 


h^ ^ 






/ V -t 




^ V 


. f 


" > /" 


-'i X 


*^ / 

A" 






38" 50' 
5'30 



Figure 2. Detail of pluvial hydrography of southwestern part of study area ('D" on fig. 1). 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



13 




38°30' 

n5»30 



Figure 3. Detail of modern hydrography of southwestern part of study urea CD' on fig. I ); showing also key 
geographic features and Locations for Gila (G-) and Rhinkhtliys (R-). 



14 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



leiiaces of the headwater stream courses ( men- 
tioned below) on the Big Smoky Valley side of 
the relatively gentle divide that separates that 
valley from Grass Valley. 

Topographic evidence indicates that these 
headwater streams at the northern end of the 
drainage basin of Big Smoky Valley (fig. 3). 
within which the large pluvial Lake Toiyabe 
existed, flowed northward into Grass Valley 
prior to a natural diversion southward, probably 
in Pleistocene time. As we noted earlier ( Hubbs 
and Miller, 1948b. p. 45). we found, northeast 
of Lake Ranch, what appears to be a stranded 
stream terrace that extends northeastward across 
the gentle divide between the present basins. Each 
of the two headwaters of the stream course in Rye 
Patch Canyon grades, at first northward toward 
a gentle pass into Grass Valley, just short of which 
each hooks backward in a hairpin turn to grade 
southward, so that flood waters discharge into 
the Toiyabe basin. One of the creeks at first flows 
northward and another, which heads near Box 
Springs, initially flows north and then northeast 
pa.st Lake Ranch (see Mount Callaghan 15- 
minute quadrangle, 1956). A dry canyon on the 
north side of each divide appears to represent the 
former, now stranded, continuation of flow 
toward Grass Valley. The evidence is clearer on 
the east side, where no 40-foot contour crosses 
the pass between the stream (the outlet of Indian 
Ranch Spring) and the Grass Valley drainage, 
between the parallel 6.520- and 6.560-foot con- 
tours. On the west side, where the stranded ter- 
race was observed, the southward diversion seems 
to have been much earlier, for the stream, where 
flowing northeastward beyond Lake Ranch, is 
rather deeply entrenched, and the north end of 
the valley has been even more downgraded, well 
below the pass altitude (between the 6.560- and 
6.6()()-foot contours). 

Rkmnant Waters and Fish Life. 

Our local inquiries and field reconnaissance of 
August 9-10. 1938. led us to believe that the only 
waters in the Lake Gilbert system that retained 



native fish until modern time are tho.se in the 
partly cut-off valley in which the headquarters of 
Grass Valley Ranch are located, west of the .south 
end of the main depression. These waters, largely 
spring-fed, are in the lower valley of Callaghan 
(Woodward) ("reek. This stream and its main, 
forked tributary. Skull Creek, rise on the east face 
of Mount Callaghan, which is assigned the al- 
titude of 10,187 feet (3,105 m. ) on the Mount 
Callaghan Quadrangle. We understand, from in- 
quiries and from examination of the topographic 
map, that the flow from these streams is ditched, 
at times of adequate flow, onto the spring-fed 
marshes, for supplementary irrigation. When the 
fish sample ( R9 ) was obtained, in 1938, a moder- 
ate quantity of water flowed, with slight to moder- 
ate current, through the wide wet meadow on 
Grass Valley Ranch, in Sec. 10, T. 21 N., R. 46 
E. The creeks named above, along with Steiner 
Creek on the eastern side of the same arm in Grass 
Valley, are depicted as permanent on the avail- 
able topographic maps (Mount Callaghan 15- 
minute and Millett I ;25(),000), but we found the 
lower courses dry in August, 1938. We think it 
highly improbable that any of these canyon 
streams, or any in the main arm of Grass Valley, 
have retained native fish. 

The only native fish that seems to have held 
out until modern times in this spring-fed meadow 
(or anywhere in the whole basin) is the strongly 
modified dace that we treat as a subspecies of 
Rliinicluhys osciiliis, under the name of R. o. 
rcliquiis (pp. 121-128). In concordance with the 
hydrographic evidence, its distinctness seems to 
suggest not only its long habitation of a single 
spring area but aLso long isolation. The only 
other main valley springs in the basin, Waiti Hot 
Springs, which lie on the valley floor east of the 
playa, in Sec. 33. T. 24 N., R. 48 E., were found 
to be far too hot for fish life. The streams that 
flow toward the valley from the higher mountains 
seem to be too impermanent and too subject to 
violent floods to have maintained fish since 
pluvial time. The apparent recent extinction of 
R. (>. reliqinis is discussed later (p. 1 23 ). 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



15 



Whether fishes other than Rliiniclitliys occurred 
in the basin of Lake Gilbert in pluvial time is 
problematical. On the one hand, the steepness 
of Cortez Canyon (dropping about 265 m. in a 
straight-line distance of only 5 km.) suggests the 
possible exclusion of species not adaptable to 
swift water. However, the great difference in 
elevation of Grass and Crescent valleys may have 
been caused by tectonic movement since Lake 
Gilbert discharged. Furthermore, the occurrence 
of chubs of the subgenus Sipluitelcs as well as of 
Rliiniclitliys in the basin of Lake Toiyabe ( Hubbs 
and Miller, 1948b, p. 45) suggests that the estab- 
lishment of those genera in the Toiyabe basin was 
most plausibly effected by a stream connection, 
discussed above, over the low pass from the basin 
of Lake Gilbert. The great differentiation of the 
Rliiniclitliys osciiliis in Grass Valley and the lesser 
modification of this species in Big Smoky Valley, - 
however, could be cited as inconsistent with this 
hypothesis, or as an indication of its early pluvial 
transfer into the Lake Toiyabe basin, followed by 
greater differentiation in the Lake Gilbert basin. 

Pluvial Lakl Diamond 

Drainage basin in the extreme southwestern 
corner of the southern part of Elko County, over 
most of Eureka County, into the southeastern part 
of Lander County, and through the long Monitor 
Valley in Nye County, all in central Nevada ( figs. 
2. 7). 

The large pluvial lake was named Diamond 
Lake by Hubbs ( 1941a, p. 65, fig. 4) and Lake 
Diamond by us (Hubbs and Miller, 1 948b, pp. 
34-35, 148, 157. figs. 10-12). This name has 
been accepted by Snyder et al. ( 1964) and Feth 
(1964). Its position es.sentially coincides with 
that of the large flat of Diamond Valley, with a 
southwestern tonguelike extension through the 
antecedent mountain gap known as Devils Gate. 



- The Rhiriichthys of Big Smoky Valley, as represented by nialerial 
from Charnock Springs, has been described as a distinct species, R. 
lariversi (Lugaski, 1972), but our study of abundant material from this 
valley has led us to regard this local form as a rather weakly differ- 
entiated subspecies of R. osculwi. 



The drainage basin, much more extensive than 
Diamond Valley, is bounded on the north by the 
watershed of Humboldt River of the Lahontan 
system; on the east by the drainage basins of 
Humboldt River, and of lakes Newark and Rail- 
road: on the .southeast, south, and southwest by 
the "Area of Sterile Basins," as treated by us 
(Hubbs and Miller. 1948b, pp. 45-51 ); and on 
the west by the watersheds of Lake Toiyabe ( not 
dealt with in this report) and Lake Gilbert (dis- 
cussed above), and by the Pine Creek division of 
Humboldt River. 

The Lake Diamond drainage basin comprises 
an elongate-elliptical, north-south oriented north- 
eastern division. Diamond Valley, which con- 
tained almost all of the ancient lake at its highest 
levels and all of the lake during much of its his- 
tory, and a larger drainage area to the west, 
through Kobeh Valley, and thence far south- 
ward. The basin in that direction comprises the 
large expanse of Monitor Valley, and, near the 
middle, also Antelope Valley, together with the 
intervening Antelope Range. The southwestern 
extension of the Lake Diamond drainage basin 
certainly embraces the watershed of Lake Diana, 
in Monitor Valley, and, doubtfully, that of Lake 
Yahoo, in the Fish Creek Range, between Dia- 
mond and Antelope valleys. Those two small 
lakes and their basins are separately treated be- 
low. 

The entire drainage basin measures 208 km. in 
greatest length and. not far north of the middle, 
85 km. in maximum width. The watershed of the 
Diamond Valley section extends 91 km. north to 
south and 33 km. west to east. Corresponding di- 
mensions for the Monitor Valley section are 163 
and 62 km. 

We estimate that Lake Diamond at its highest 
(outlet) level was 70 km. in greatest straight-line 
length and 19 km. in maximum width. At the 
somewhat lower stand, which we have also 
mapped ( as a dashed line ) , these dimensions were 
55 X 18 km. We compute the maximum area of 
the lake as 1,002 sq. km., constituting 15 per- 
cent of the 6,758 sq. km. of the entire drainage 



16 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



basin, including that of lakes Diana and ^ ahoo. 
At the lower level these figures are 792 sq. km.. 
I 2 percent of the basin. 

Inasmuch as the altitude of most of the periph- 
eral rim of the Lake Diamond drainage basin ex- 
ceeds 2,500 ni. and the basin lies in the latitudinal 
band of high pluvial precipitation and of high 
ratio between lake and drainage-basin area, it is 
.surprising that the ratio for this lake is at most 
only I 5 percent. This circumstance we interpret 
as corroborative evidence that the lake discharged, 
as we have indicated ( Hubbs and Miller, 1948b, 
pp. .M-.^6. figs. 10-12). 

With the exceptions certainly of Lake Diana, 
probably of Lake Yahoo, and possibly of Lake 
Snyder (see discussions below), no pluvial lakes 
can be postulated to have discharged into the 
drainage basin of Lake Diamond. 

Deposits in the northern part of Kobeh Valley 
that were mapped by Merriam and Anderson 
(1942) as Quaternary, including lake beds, are 
too high to represent the western arm of Lake 
Diamond as we interpret it, but possibly represent 
the remains of a lake of some earlier period. 

The ground waters of Diamond Valley have 
been described by Eakin ( 1962) who also mapped 
the shoreline of the ancient lake. 

Shoreline and Discharge. 

Reconnaissance determinations in I9.^S of ter- 
race elevations on the west side of Diamond Val- 
ley, toward the north, with the use of a Paulin 
altimeter, seem to confirm the idea that Lake 
Diamond discharged through Railroad Canyon. 
The altimeter readings seem to place the maxi- 
mum altitude of the lake somewhat above the 
6,()()0-foot ( l,S29-m.) contour as shown on the 
Winnemucca. Millett, Elko, and Ely 1:250,000 
maps and on the 6 pertinent 15-minute quad- 
rangles (Mineral Hill 19.-^7. Railroad Pass 1959. 
Garden Valley 1959. Diamond Springs 1957, 
Whistler Mtn. 1956. Eureka 1953). Since the 
6.0()0-foot contours are very closely approxi- 
mated along Railroad Canyon, but do not con- 
join there, it is concluded that the spillase down- 



graded the can_\on and thus lowered the level of 
the lake while it continued to overflow. Weakness 
of terraces higher than the lip of the canyon seems 
to confirm this view. The present altitude of Rail- 
road Pass is given on the Railroad Pass Quadran- 
gle as 5,895 feet ( 1,797 m. ). Since some of the 
lateral terraces were judged to be at an elevation 
of about 5,900 feet ( see below ), it is thought that 
the lake stood at aboLit this height for some time. 
Our maps (figs. 2. 7) show lake borders at ap- 
proximately 5,900 feet ( 1.798 m.), for the last 
outlet stage, as well as at about 6,000 feet ( 1 ,829 
m.). to approximate the assumed initial outlet 
stage. Because the lake impinged against the 
steep west and east sides of the valley, these two 
lake levels are in general very closely approxi- 
mated, except at the south end where the valley 
slope is very gentle. 

The close approach of the same 6,0()0-foot con- 
tours through the inlet gap. Devils Gate, as well 
as in the outlet gap. Railroad Canyon, sug- 
gests that a tongue of the lake, at the highest 
level, extended westward through this trench 
slightly onto the broad desert flat beyond. Ac- 
cording to the Whistler Mtn. 15-minute Quad- 
rangle, checked by the planetable survey of 1956, 
the 6,02()-foot (1,8.^5-m.) interpolated contour 
extended 10.7 km. west of Devils Gate, in the 
mouth of Antekipe Valley east of Lone Mountain. 
Assuming the outlet level at 6.015 feet ( 1.8.^.3 m.: 
see below ). the tongue of the lake is estimated to 
have extended about 9.5 km. west of the Gate. 
The exact extent of the tongue is a bit uncertain, 
for the slightly earlier Millett 1:250.000 map 
shows the 6,()00-foot contour extending 12.5 km. 
beyond the Gate, to a point southeast of Lone 
Mountain. 

The deep trench at Devils Gate, west of Eureka, 
appears to be an antecedent water gap in the 
north-soLUh file of mountains flanking Diamond 
Valley on the west, and the gap appears to date 
from some previous drainage system, it has 
obviously served, at all but the very highest stage 
of Lake Diamond, as the channel through which 
pluvial Monitor River flowed to reach Lake Dia- 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



17 



mond. A figure in our former report ( Hubbs and 
Miller. 1948b. fig. 12) made from a photograpli. 
shows what may be a stream terrace on the south 
side just above the west end of this inlet pass. 

On the west side near the north end of the lake, 
toward the south end of an island (or peninsula), 
a strong terrace (shown as fig. 10 of our 1948 
report) was found to be close to the level of the 
outlet pass (Railroad Canyon), and traces of 
slightly higher shorelines were seen here. At Big 
Shipley Spring (Sadler Ranch. T. 24 N.. R. 52 
E. ), altimeter readings taken rapidly and checked 
before and after against Bench Mark H 10 1936 
(assigned elevation 3.836 feet), indicated the 
sharpest terrace at 6,020 feet (1.835 m.) and 
others at 5.915?, 5,965. 6,100?. and 6.075? feet. 
A second transect, taken farther south in the 
same township, midway between the Sadler and 
Bailey ranches, showed a broad terrace at about 
5,895 feet ( 1,797 m. ). well below the very strong 
terrace north of the middle of T. 23 N., R. 52 E., 
on the point just north of the Romano Ranch. 
That terrace was checked at 5,945 feet (1,812 
m. ) and a lesser terrace with the highest level of 
waterworn cobbles was tallied at 6,015 feet 
( 1,833 m.). Several other terraces were traced, 
some well above the main outlet level, all around 
the north end. and on the faces of projecting 
points, which must have been exposed to waves, 
along the west side of the ancient lake. A sharp 
but rather low terrace rounds the point on the 
north side of the eastern end of Devils Gate, ex- 
tending onto the desert flat, but this may have 
been a stream terrace. On the flat southeastern 
side of the basin, shorelines fade out, as is to be 
expected on such areas, where marsh vegetation 
no doubt inhibited shoreline erosion. Except 
locally, terraces are weak on the east side of the 
flat lake bed in Diamond Valley. Receding lake 
levels are suggested by dunes seen near the east 
edge of the alkali flat in the southern part of T. 
25 N.. R. 54 E. and by curved elevations shown 
on the topographic maps. These maps, however, 
show little in the way of .shoreline features, pre- 
sumably because the lake, during all but the latest 



stages of recession, impinged on the generally 
steep bajada. If the maximum altitude of the lake 
be regarded as 6.015 feet, and the altitude of the 
playa is taken as 5,767 feet, the depth of the lake 
above the playa level would have been 248 feet 
(76 m.). At the lower level mapped, the depth 
would have been 45 m. 

Remnant Waters and Fish Life. 

Diamond Valley. Most of the spring waters 
in the Lake Diamond drainage basin lie in Dia- 
mond Valley, which we reconnoitered in August, 
1938. Here we found numerous spring waters, 
some profuse, mostly within but toward the edge 
of the flat shown on .some maps as "Alkali 
Desert." These springs are more numerous on the 
western side of the valley, along the fault line at 
the base of the steep escarpment of the Sulphur 
Spring Range, than on the eastern side, along the 
foot of Diamond Mountain. We probably saw a 
sample of most of the springs on both sides. We 
treat the springs counterclockwise from the 
north : 

"Lake Dou-pah-gate"' was shown on the Land 
Office map of Nevada ( 1 930 ) as much too large. 
This spring pool, in the northwest corner of T. 25 
N.. R. 53 E., was found to be only about 30 m. 
in diameter, very deep and chalky blue. It was 
reported to contain "bullheads" (presumably 
Ictulunis sp. ). but no minnows. Careful observa- 
tion around the edge and in the outlet slough and 
ditches disclosed no fish. This was probably one 
of two springs shown near Flynn Ranch on the 
Mineral Hill 15-minute Quadrangle, which also 
shows Josephine Spring one mile to the south. 

Big Shipley Spring (of 1938), later mapped as 
Shipley Hot Spring, at Sadler Ranch ( Location 
R8, figs. 3, 8). formed a large spring pond (fig. 
21 ), or small lake, about 3 m. deep, of clear, 
warm, .somewhat sulphurous water, with consider- 
able vegetation. The cluster of head springs 
registered 39 C. The outlet was reported by a 
Biological Survey rodent-control agent then work- 
ing in the area to extend about 5 miles (8 km.) 
onto the alkali flat, where some water accumu- 



11 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



lates. The collection comprised goldfish and a 
local form, presumed to be native, of the speckled 
dace, Rhinichlhys osciilii\ (p. 115). 

A spring just south of Big Shipley Spring was 
reported by the rodent-control agent to ccMitain 
similar fish. 

A rather large, very soft-lx>ttomed spring hole 
in the yard of Bailey Ranch, in the southwest cor- 
ner of Sec. 36, T. 24 N.. R. 53 E.. 3.2 km. south 
of Big Shipley Spring, was reported by the same 
informer to contain "chubs"' and rainbow trout. 
Unfortunately we failed to check on this report, 
for the "chubs" may have repre.sented a third 
population of Gila hicolov in the Lake Diamond 
dramage system. Or, this ranch-yard pool may 
have been stocked with chubs from Sulphur 
Spring or the Birch Ranch, for mosquito control 
or other reason. 

Springs shown on the Ciarden Valley Quad- 
rangle in T. 23 N.. R. 52 E. between the Bailey 
and Romano ranches, at Romano Ranch, and for 
3.2 km. farther south, the southernmost mapped 
as TliIc Dam Spring, all obviously along the .same 
fault Ime, were also not examined, but local testi- 
mony gave no indication that they contained fish. 

Sulphur Spring (Location G6. figs. 3. S). 14 
km. by road south of Sadler Ranch, and now 
mapped as in Sec. 36. T. 23 N., R. 52 E. on the 
Ciarden Valley I 5-minute Quadrangle, formed a 
partly artificial pool of clear water about 10 m. 
wide (described in some detail on page 154). It 
harbored a form that we interpret as a distinctive 
race of (Jila hicolor obesn (Girard), one of the 
two we found in the whole drainage system. That 
this spring was probably long known to travellers 
is suggested by the long-used name of Sulphur 
Spring Mountain, that rims Diamond Valley on 
the west side. 

The Ciarden Valley 1 5-minute Quadrangle 
shows a cluster of other springs within about 1 
km. of Sulphur Spring. It is possible, though we 
think unlikely, that .some of these springs and 
others mentioned above, all along the same fault 
line, may contain a remnant oi native fish. There 
appear to be no prospects farther south on the 



west side of Diamond Valley, nor around the 
south rim. 

A .spring at Maggini Ranch, on the east side of 
the valley in Sec. 27, T. 23 N., R. 54 E., was 
found to be very small and fishless. The same 
almost surely applies to the two springs shown on 
the Diamond Springs Quadrangle in the same 
township between the Maggini and Thomp.son 
ranches. Close to one of the.se two ranches there 
is a small subterranean body of clear water known 
as Emerald Lake, in Emerald Lake Cave. Local 
claims of fish having been seen in this under- 
ground water have not been verified by Robert H. 
Soulages, biology student and active speleologist 
(personal communication. 1970), who will con- 
tinue to kxik for fish here. 

Two large springs on Birch Ranch (in 1938 
the .lorge Jacobsen Ranch, later the Thompson 
Ranch), just off the base of Diamond Mountain, 
in Sec. 3 of the same township, yielded both 
Rhiniclithys osciiliis (p. 115) and Gila hicolor (p. 
154); mapped as Location G5 and R7 (figs. 3. 8). 
This is one of the two localities in all the basins 
under treatment that have retained both species. 
As noted later (p. 115), Mr. .lacobsen. who in 
1938 had been on the ranch for 35 years, believed 
that the dace and chubs were native, and stated, 
in agreement with our findings, that fish occur 
on the east side of Diamond Valley only on his 
ranch. 

These spring waters are apparently the south- 
ernmost o\ consec|uence on the east side of the 
great alkali flat. Proceeding northward we 
checked the ranches in an effort to locate other 
fish-inhabited waters (the narrow western slope 
of the Diamond Mountains offered no expecta- 
tions). There were ranches at 1.4 and 10.6 km. 
along the almost straight road between Birch 
Ranch and Railroad Canyon, with no indication 
of sizable springs. Diamond Springs are named on 
the Diamond Springs Quadrangle at the site of 
the first ranch, and others are shown about a mile 
farther north. No springs were seen around the 
north end of the old lake bed. and none is shown 
there, save for a few small mountain springs, on 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



19 



the Mineral Hill and Railroad Pass 15-minute 
quadrangles. 

Monitor Valley. Exploration and inquiries 
in 1938 led us to conclude that permanent surface 
water and fish, other than stocked trout, are con- 
fined to two spring areas, namely Coils Creek 
and Potts Ranch, in the wide expanse ( see above ) 
of Monitor Valley. Both areas were sampled. In 
1972, fish were found in the discharge of hot 
springs about Dianas Punch Bowl, south of the 
Potts Ranch hot spring. 

At the Three Bar Ranch on the course of Coils 
Creek (Location R6. fig. 3). near the north end 
of the basin ( shown on the Roberts Creek 1 5- 
minute Quadrangle in Sec. 4, T. 22 N., R. 49 E.), 
a spring pool in grassy meadows yielded a large 
sample of one of the innumerable local forms of 
speckled dace. Rhinichthys o.sciiliis (p. 114). 

In addition to Coils Creek, two mountain 
streams, Roberts and Ferguson creeks, are 
mapped as permanent, but each goes under- 
ground on a large alluvial bajada. We long 
thought that neither would seem likely to have 
retained fish. When examined on August 14, 
1938, the stream at Roberts Creek Ranch was 
found to be very small, and it was locally reported 
to contain only stocked trout. However, one in- 
formant, biology student Robert H. Soulages (per- 
sonal coinmunication, 1970). has stated that he 
has seen small fish farther up Roberts Creek that 
he felt strongly were not trout. The locality is 
where the stream, near its forks, flows in a rather 
flat valley, as well as in a canyon (as shown in an 
aerial photograph supplied by Mr. Soulages). 
These fish may well have been speckled dace. 
Ferguson Creek was not examined, but the topo- 
graphic maps raise little expectation that it con- 
tains native fish. 

What we then thought to be Twin Springs on 
Bartine Ranch, on the flat area near the junction 
of Monitor, Antelope, and Kobeh valleys, were 
said by the rancher on August 15, 1938, to be of 
artesian origin and to contain no fish, and none 
were seen by us in the ditch and tank by the road. 
However, the Bartine Ranch 15-minute Quad- 



rangle shows artesian wells at the ranch, and Twin 
Springs, Cold Spring, and Warm Spring, nearly 
2 miles west, and other springs to north and south. 
Because springs in this location seemed to hold 
some chance of having retained native fish, those 
just named were checked by the Miller family on 
July 3, 1970. They were found to be of limited 
flow, without trace of fish, and to lack features 
associated with the retention of fish. 

The other significant spring area in Monitor 
Valley for which we found evidence was at the 
hot springs ( Locations R5 and R5A, fig. 3 ), near 
the south end of the axial stream course of the 
valley, now the largely dry southern branch of 
Stoneberger Creek. 

One of the hot springs in this area was found 
on Potts Ranch (in Nye County. 9 km. south of 
Lander County), earlier known as Wilson Ranch. 
Here, clear water was found on Augu.st 16, 1938, 
to issue at 42 C. In the outflow, in a gras.sy 
meadow, where the water had cooled to 32-34 
C. another local form of Rhinichthys osvuhis 
rdbusiiis (Rutter) was taken (p. I 13). That the 
dace is endemic in this spring is indicated by the 
lack of any contrary evidence, by the rather 
copious water supply, by the location of the spring 
to one side of the main flood channel of the valley, 
and by the evidence of long continuity of spring 
flow furnished by the extensive travertine de- 
posits beside which the water issues. The habitat 
and the fish population are described and dis- 
cussed in some detail (p. 1 14. fig. 20). 

During our reconnaissance in 1938, a local in- 
formant, a Forest Ranger named Crane, expressed 
the opinion, with which rancher George Potts 
agreed, that there is no reasonable chance that 
any fish other than stocked trout exist in any other 
waters of Monitor Valley (or in the valleys of 
Ralston and Stone Cabin creeks to the south). 
Nor did we find, through travel, inquiry, and ex- 
amination of available maps, any other evidence 
of native fish life in Monitor Valley, or elsewhere 
in the entire western, intermittent drainage of 
Diamond Valley, except in the meadow area of 
Coils Creek and possibly in Roberts Creek (both 
discussed above). 



20 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Another hot spring issues, close to the course of 
pluvial Monitor River, in a meadow area of 
moderate size about 5 km. south-southwest of Hot 
Springs on Potts Ranch. This meadow was thought 
in 1938 to be fishless. but may have contained 
the ancestors of the fish found in 1972 in the 
hot-spring discharge discussed below. Farther on, 
1.2 icm. in the .same direction, is another large, 
fishless hot spring known as Dianas Punch Bowl, 
which, strangely, forms a deep pool of clear water, 
which on August 16. I9.^(S, was 7.5 m. in diam- 
eter, with a surface about 7 m. below the summit 
of a broad travertine dome measured by 1-foot 
altimeter as about 15 m. high (formerly the level 
was higher and now inactive holes on the sloping 
sides indicate former discharges). 

Other hot springs ( Location R5A ) are shown 
on the Dianas Punch Bowl 1 5-minute Quadrangle 
(I960) as arising beside the road immediately 
southwest of Dianas Piuich Bowl, and as forming 
the head of flow of the south branch of Stone- 
berger Creek. These springs we failed to see when 
the area was explored on August 16, 193(S, and 
may well have then had a very limited discharge, 
in the meadow area that was then observed. How- 
ever, in 1972 these springs yielded a stream flow 
for a short distance, in an apparently recently 
constructed ditch, and were found to contain a 
population of Rhinichthys nscaliis ( Robert E. 
Brown, personal comniunicatiiMi ). The fish are 
in many respects like those of Potts Ranch Hot 
Springs, but sufficiently different in some details 
to suggest some degree of separation of habitat 
and of fi.sh. In February, 1972. the main, west- 
ernmost of the springs in the ditch issued at 45 C. 
After it had turned northward the ditchlet perco- 
lated through a porous travertine deposit. Tem- 
peratures where fish were collected 100 m. above 
the barrier had cooled to 37.5 , and about 200 ni. 
below the barrier to 37.0 . The flow below the 
barrier was about 0.3 cubic feet per second. 

Slightly farther south, at approximately 39 
00' N. lat.. Box Springs, with a small fishless out- 
flow, was found in 193K to maintain a small 
meadow on the west face of an OLitlvinc hill of 



the Antelope Mountains. This lava hill, along 
with an alluvial cone from Mill Canyon (in the 
Toquima Range), dams the axial drainage of 
Monitor Valley to form an ephemeral lake, with- 
out outlet, the site of small pluvial Lake Diana (p. 
21 ). At the time of our 1938 visit, this lake con- 
tained much water, following rains, but ordinarily 
it has no surface water and is of course fishless. 

Except for the waters just described, in and 
near the course of ancient Monitor River, the 
surface waters of Monitor Valley are confined to 
very limited spring-fed sections in mountain 
canyons. Such habitats are almost always devoid 
of native fish life. Local testimony agreed with 
the expectation that no mountain streams in the 
valley are of sufficiently low gradient to have 
maintained native fish. 

Evidence was also obtained in 1938 of the lack 
of native fish in the Antelope Valley arm of the 
southwestern part of the flood-water drainage sys- 
tem of Diamond Valley. Forest Ranger Crane 
and rancher Potts referred us to the rancher at 
Cerutti Ranch in Copenhagen Canyon in the 
Antelope Valley drainage for information on pos- 
sible, though they thought improbable, native fish 
in Antelope Valley. That rancher was sure that 
no fish occur in the streamlet on his ranch (prob- 
ably the one labelled Martin Ranch on Horse 
Heaven Mts. 1 5-minute Quadrangle), or else- 
where in Antelope Valley, wherein we saw none. 
We found Hot Spring in the same valley, shown 
on the Antelope Peak 1 5-minute Quadrangle in 
Sec. 28, T. 1 8 N., R. 50 E.. to be too hot for fish. 
Although some streams are mapped as permanent 
in the adjacent nioLintains. we think it highly im- 
probable that any native fish exist in the drainage 
system of Antelope Valley. We were intrigued 
with the name of "Fish Creek Well" in a canyon 
on the east side of the valley close to the probable 
course of the ancient outlet of Lake Yahoo, but 
the location seems most improbable for a native- 
fish habitat. Probably the well was named for 
Fish Creek Ranch and may have been drilled by 
Mr. Fenstermaker. the pioneer at that ranch 
across the Fish Creek Ranue. Fenstermaker 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



21 



Wash, in Antelope Valley, is not far south of Fish 
Creek Well. 

Pluvial Lake Diana 

Drainage basin comprising the upper, southern 
part of Monitor Valley, near middle of north 
border of Nye County, central Nevada ( fig. 2 ) . 

This very small lake, almost surely of pluvial 
origin and represented at present by the ephemeral 
lake discussed above, was named by Snyder et al. 
(1964). It was not discussed by us in our 1948 
report, but was shown on our map of pluvial 
waters. In pluvial as well as modern time, as 
noted above, the lake appears to have been 
dammed by an alluvial cone, from the Toquima 
Mountains, impinging on a lava hill that juts into 
the valley from the Antelope Mountains. 

The north margin of the drainage basin cuts 
across the Monitor Valley arm of the Lake Dia- 
mond watershed; to the cast lies the Little Fish 
Lakes arm of the Lake Railroad system; to the 
southeast, south, and southwest occur tributaries 
of what we have called the "Area of Sterile 
Basins" (Hubbs and Miller. 1948b, pp. 45-51 ); 
and to the west, the drainage basin of Lake 
Toiyabe. 

The roughly oval drainage is 58 km. in maxi- 
mum ( north-south) length and 31 km. in greatest 
width. Because of its position in the stream 
course down the axis of Monitor Valley, the 
pluvial lake presumably had, as Snyder el al. 
depicted it, approximately the form and dimen- 
sions of the present ephemeral lake, 9 km. long 
and 2 km. in greatest width (near the southern, 
inlet end). We estimate the area of Lake Diana 
as only 12 sq. km., I percent of the area (1,339 
sq. km. ) of the drainage basin. 

Since this obviously integral part of the Lake 
Diamond hydrographic basin is bordered on each 
side by high mountains, mostly more than 2,700 
m. in altitude, it presumably contributed much of 
the flowage in pluvial Monitor River, the main 
affluent to Lake Diamond. At present, its per- 
manent waters are confined to the mountains and. 



we assume, are devoid of native fish life. We did 
find, however, as noted above, a population of 
speckled dace, Rliinichlliys osciilus, in the valley- 
spring water of Potts Ranch, and about Dianas 
Punch Bowl, which lies just below ( north of) the 
location of Lake Diana. 

Pluvial Lake Yahoo 

Drainage basin in southeastern Eureka County, 
central Nevada (figs. 2, 7). 

This lake was shown on our 1948 map, but was 
neither named nor discussed. It was mapped and 
indicated as a Pleistocene lake by Snyder et al. 
{ 1964) and by Morrison ( 1965, fig. 1 ), but was 
not named by them. Snyder et al. listed the val- 
ley as "Yahoo" but we note some discrepancy in 
the name. The modern remnant bears the non- 
committal name Dry Lake on the General Land 
Office map ( 1930). and the Bellevue Peak 15- 
minute Quadrangle ( 1956 ) names Dry Lake Well 
along the margin of the playa. The name Yahoo 
was, we suppose, derived from the name of the 
canyon and creek that heads north beyond a low 
mountain ridge, although the apparent fault block 
basin opens southward into Spring Valley (tribu- 
tary to Antelope Valley) along the course that 
we assume was taken by the ancient outlet of the 
lake. On the Millett 1:250,000 map the name 
given is Yahoe Creek, whereas the Whistler Mtn. 
1 5-minute Quadrangle gives Yahoo Canyon. We 
suggest that the ancient lake be called Lake 
Yahoo, though it was of minimal size. 

This minute drainage basin probably was in- 
cluded within, and certainly is entirely surrounded 
by the watershed of Lake Diamond (pp. 15-20), 
though it almost abuts on the Newark-Diamond 
divide. It is presumably a fault-block depression 
in the mountains, with the lowest pass, on the 
south side, above the 7.320-foot (2.231-m. ). but 
below the 7,360-foot, contour (as shown on the 
Bellevue Peak 1 5-minute Quadrangle). As we 
have drawn the basin and lake borders after 
Snyder et al.. we measure the length and breadth 
of the basin as 12 and 6 km., and the length and 



22 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



breadth of the pluvial lake as 3 and 2.5 km. We 
eompute the area of the lake as 7 sq. km., eon- 
stitiiting I 6 pereent of the area. 44 sq. km., of the 
drainage basin. 

The topographie maps indieate that an inter- 
mittent playa pond exists within the old lake bed. 
Snyder ct al.. on the basis of field work by Hard- 
man, showed an outlet extending southward, then 
westward, to join the axial stream eourse of Ante- 
lope Valley, of the Lake Diamond system. We 
have no reason to question that eonelusion, in 
view of the height of the surrounding mountains 
and the drainage relations of the adjacent basins. 

We have not examined the basin, which we 
suppose has only intermittent surface water and 
no fish life. 

Pluvial Lakh Ni:\vark 

Drainage basin in far-western White Pine 
County, the southeastern corner of Eureka 
County, and extreme northern Nye County, cen- 
tral Nevada (figs. 2. 7). 

This lake, the main one in the area undei' treat- 
ment, was named anti briefly discussed by us 
(Hubbs and Miller. iy4Sb. pp. .^.^-.^4). This 
designation has been accepted by Snydei' ct al. 
{ 1964). Feth ( 1964). and others. 

The drainage basin is bounded on the north by 
Huntington Creek of the Humboldt River system 
(its inferred high-level outlet): on its northeast 
sector for only 6 km. ( straight-line measurement ) 
by a minor tributary of Lake Franklin: on the east 
mostly by the once possibly tributary basin of 
Lake Hubbs, and southwaid. for a straight-line 
(.listance of 16 km., by the Lake Jake watershed; 
on the southeast and south by the drainage basins 
of Lake Railroad and of Lake Limai Crater and 
Lake Snyder, both perhaps tributary to Lake 
Railroad; and on the southwest and west, for 
about 15 km. each, by the Hot Creek and Little 
Fish Lake sections of the Lake Railroad basin: 
farther north on the west side, by the Lake Dia- 
moni.1 hydrographic system. 

The extensive area that drained in the pluvial 



period into Lake Newark comprised not only a 
northeastern part. Newark Valley proper, with a 
southern extension east of the Pancake Range, 
but also a southwestern arm. Fish Creek Valley, 
which is also connnonly mapped as the northern 
part of Little Smoky Valley; the southern part of 
that arm is shown as extending not only across 
the southernmost part of the Newark drainage 
basin but also to and across the basin of pluvial 
Lake Snyder (pp. }!-?>><}. Therefore, Fish Creek 
Valley seems to be regarded as a part of either 
Newark Valley or Little Smoky Valley. Even in 
historic time, Fksh Creek Valley has drained oc- 
casionally into Newark Valley, but the valley is 
partly shut off from the Pancake Range by a 
mountainoLrs |irominence known as Black Point, 
which constricts the graben. Pancake Range sepa- 
rates the two southern extensions of the Newark 
drainage .system. 

Lake Newark occupied what is now an elon- 
gated playa, extending, as do the marginal moun- 
tains, north-south in Newark Valley and north- 
east-southwest in Little Smoky Valley and the 
connecting area. The extreme length of the total 
drainage basin is 132 km.; the greatest width, at 
middle. 55 km. The least width near the middle 
of the noilhern and southern parts, respectively, 
is 17 and 22 km. The lake area roughly conforms 
with that of the basin, but is displaced northward. 

At the assumed outlet level the greatest straight- 
line length of the lake wmild have been S8 km. 
and the maximum width, in the southern part of 
the northern area. 20 km. The least widths, near 
the middle of the northern and southern .sections, 
would have been 9 and 5 km., respectively; the 
greatest widths, toward the north and south ends, 
1 4 and 1 7 km. 

The area of the lake, at the discharge level, 
would have been 1.146 sq. km., or 32 percent 
of the area ( 3.587 sq. km. ) of the drainage basin. 
When, if ever. Lake Hubbs discharged into Lake 
Newark (see p. 27). the drainage would have 
been increased to 5.264 sl|. km. and the area of 
the lake would have constituted only 22 percent 
of the combined drainaiie area. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



23 



At the lower, unquestionable lake level shown 
on two of our maps ( figs. 2,7) as a dashed line, 
the lake dimensions would have been as follows: 
greatest straight-line length, 76 km.; maximum 
width. 18 km.; least width, north and south, 8 
and 3 km.; area, 792 sq. km., or 22 percent of 
the drainage-basin area (exclusive of the Lake 
Hubbs basin). 

The depth of Lake Newark above the level of 
the present alkali flat is estimated on the basis of 
the following numbers. The lowest assigned alti- 
tude of the alkali flat, which appears on the Buck 
Mountain, Cold Creek Ranch, and Eureka 15- 
minute quadrangles, is 5,833 feet (1,778 m. ). 
The outlet-level contour as noted below is treated 
as slightly above the 6,200-foot contour and the 
lower level is treated as midway between the 
6,040- and 6,080-foot contours. Assuming figures 
of 6,210 and 6,060 feet, the lake depths (above 
the present playa) are computed as 377 feet (115 
ni. ) and 227 feet ( 69 ni. ). Snyder et cil. gave an 
estimate of 285 feet (87 m.). The lower level 
we have drawn is probably over conservative. For 
example, as noted below, a definitive level just 
above Fish Creek Springs seems to be only about 
45 m. below the outlet level. We regard the 
lower level drawn as uncontroversial and as rep- 
resenting a long-period steady state. This was 
the level accepted by Rush and Everett (1966. 
p. 10). 

The large storage of water, on either lake- 
level assumption, is consistent not only with the 
position of the lake in the latitude of apparently 
heavy pluvial precipitation, but also with the 
height of the marginal mountains, especially the 
ranges on the abrupt west side (from north to 
south, the Diamond, Fish Creek, and Antelope 
ranges), which form a ridge almost uninter- 
ruptedly exceeding 2,500 m. in altitude. 

The ground waters of Newark Valley have 
been described by Eakin ( I960). 

Shoreline and Discharge. 

There is a possibility that Lake Hubbs attained 
an early pluvial discharge into Lake Newark ( p. 



27 ) and a remote possibility that pluvial Lake 
Snyder overflowed into Newark Valley (p. 38). 
That Lake Newark discharged into Huntington 
Creek and hence through Humboldt River into 
Lake Lahontan was claimed by us ( Hubbs and 
Miller, 1948b, p. 33 and map). Our field notes 
(C.L.H.) of September I 1, 1934. seem definite: 

Very clear wave-cul terraces were observed 
around the eastern side ol Newark Valley. 
They were especially sharp and numerous 
around the north end ol the valley, where they 
cut like steps into the evenly sloping valley 
sides. They reached just to the height of the 
ridge separating Newark Valley from Hunt- 
ington Valley, indicating an outlet into the 
South Fork of Huniholdt River. 

Snyder et <//., on the contrary, stated that Lake 
Newark did not spill, and C. T. Snyder (personal 
communication ) has indicated that he was not 
able to confirm such a discharge, either by field 
reconnaissance or by examination of aerial photo- 
graphs, and that the shoreline he identified rounds 
the north end of the valley below the divide. Pend- 
ing a more detailed survey, we maintain the view 
that Lake Newark did discharge during some 
pluvial, perhaps early pluvial, time. A prime basis 
for this assumption is our finding that the native 
fish of the Lake Newark basin, constituting in our 
judgment two local subspecies, are most closely 
related to Gila hicolor ohcsii. which is character- 
istic of the Lahontan hydrographic system. The 
degree of differentiation suggests a longer period 
of isolation than has befallen the Lake Diamond 
populations of Gila hicolor (and Rhiiiichtliys 
osciili(.s). We may have misjudged the height of 
the pass and may have interpreted a shoreline 
along an enclosed contour as being at the outlet 
level. However, shoreline features actually at the 
outlet level may well have become eroded beyond 
recognition during the probably long time since 
they were formed. 

Our observations lead us to conclude that the 
highest outlet level of Lake Newark lay slightly 
above the 6,200-foot ( 1 .890-m. ) contour: at two 
points on the Cold Creek Ranch 15-minute Quad- 
rangle, parallel 6.200-foot contours barely miss 



24 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




Figure 4. Shoreline features (bars and spits, pointed out by arrows) mapped along the 5, '^'20-. d. ()()()-, and (i,U40- 
toot ( l,S()4-. l.S2y-. and l,S41-ni.) eontours in the eastern emhayment. near middle, ot pkisial lake Newark, west- 
ern White Pine County. Nevada. These contours are .S7, 167, and 207 feet (27, 51, and 7.i m.) above the lowest 
stated altitude on the alkali flat of Newark Lake. Superimposed on a southeastern p.irt of United States Geological 
Survey Buck Mountain Quadrangle ( 1^)57). Dashed line at 6.()60-foot ( l,S47-ni.) altitude is regarded as representing 
a minimal lake level, presumed lo he late pluvial; heavy solid line at (i,2()()-foot ( K.S^O-m.) altitude is regarded as an 
earlier-pluvial outlet level. Beck Pass m upper-right corner represents the possible early-pluvial outlet course of Lake 
Hubbs. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



25 



touching along the northern sill of the basin, 
where the drainage basin of Latce Newark con- 
tacts that of Huntington Valley of the Humboldt 
River watershed. We have, therefore, drawn the 
margin of the lake at maximum stage close to 
that contour on the Buck Mountain. Cockalorum 
Wash, Cold Creek Ranch. Diamond Springs, 
Eureka. Moody Peak, Pancake Summit, and 
Pinto Summit 15-minute quadrangles, and have 
had these transferred onto the 1:250,000 maps. 
In addition, we have drawn a second lake bound- 
ary, midway between the 6.040- and 6,080-foot 
(1,841-m. and 1,853-m.) contours, to represent 
a level clearly shown by the shoreline terraces 
seen and by strikingly bold longshore bars de- 
lineated along the 6,040-foot line in the eastern 
bulge near the middle of the lake, on the Buck 
Mountain 15-minute Quadrangle. This map also 
.shows clear-cut recessional .shoreline features 
(bars and spits) on the 6,000-foot (1,829-m.) 
and 5.920-foot (1,804 m.) contours (fig. 4). 
Dunes close to the 6,000-foot contour along the 
west side of a small flat just south of Fish Creek 
Springs (see below), at the foot of the bajada 
from Fish Creek Range, probably are also near 
an old shoreline of long standing. 

Remnant Waters and Fish Life. 

The most copious valley waters in the Newark 
drainage basin consist of Fish Creek and its head- 
water springs on Fish Creek Ranch in Fish Creek 
(Little Smoky) Valley, and several springs mar- 
gining the flat bed of Newark Valley. 

The profuse Fish Creek Springs ( Rush and 
Everett, 1966, pp. 14, 25, 29). the .sole habhat 
of Gihi bicolor eiichiht (pp. 168-169), rise on the 
old lake bed near the western edge of its south- 
end expansion, about 5 km. west of Fish Creek 
Ranch headquarters, in the NW. '-4 Sec. 8, T. 16 
N., R. 53 E. (as is shown on the Bellevue Peak 
and Pinto Summit 15-minute quadrangles). In 
1938 there was one main spring north of two in 
tandem, with outlets soon joining in a meadow 
to form Fish Creek above the ranch headquarters. 
The more recent topographic maps show the 



spring source extensively draining into irrigation 
ditches. The spring water may originally have 
maintained Fish Creek for a considerable distance, 
but by 1938 or earlier it was usually consumed on 
Fish Creek Ranch, and was dry at the road cross- 
ing 7 km. below (east of) the springs on August 
17, 1938 (well after the 1934 drought). Isador 
Sara, the Basque who had long operated Fish 
Creek Ranch, testified when interviewed on that 
date, that about 25 years previously water from 
Fish Creek reached Newark Valley. It did so 
when he ditched the creek on the valley flat in an 
effort to irrigate some lower fields ( in the winter 
this was done to prevent the freezing of the fields ) . 
As a result, wagons mired along the old road. 
Ordinarily, he said ( as of 1938 ). the spring water 
when not u.sed for irrigation sank into a rather 
long .shallow "lake" on the flat. 

Mr. Sara held that the chubs of Fish Creek 
Springs are without doubt native, as indeed the 
name of the creek and the springs indicates. He 
know of no other native fish in the general vicin- 
ity, except those in springs about "Fish Lake" 
(obviously referring to the chubs, Gila bicolor 
subspecies, of Little Fish Lakes in a tributary 
valley of the Lake Railroad system, separated 
from Fish Creek Valley by a low divide ) . Pre- 
sumably no waters in Fish Creek Valley other 
than those of Fish Creek Springs have retained 
native fish. 

Dunes on the valley flat just off the alluvial 
slope a short distance to the southward, and other 
evidence, indicate that Fish Creek Springs arise 
just within a definitive level of pluvial Lake 
Newark, but about 45 m. below the hypothesized 
outlet level. Although Fish Creek Valley has dis- 
charged within historic time into Newark Valley, 
there is no reason to suspect that the fish-inhabited 
waters of Fish Creek and Newark valleys have 
had any postpluvial surface connection. Since the 
pluvial period, any surface discharge from Fi.sh 
Creek Valley has presumably sunk into the cen- 
tral part of the old lake bed in Newark Valley, 
short of any connection with discharges from the 
fish-inhabited springs of Newark Valley. The 



26 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



differentiation of Gila hicolor iicwdrkcnsis and 
G. b. cucliilu. the endemic fisiies of the Lake 
Newarlv system, is consistent with this view. 

Springs that margin the ancient lal<e bed ap- 
parently provide the only habitats in Newark 
Valley proper that have retained native fish. 
These fish comprise a subspecies. Gila hicolor 
ncuiirken.sis (pp. 1 5fi- 1 3S ). which we differentiate 
not only from G. h. ohiwci of the Lahontan system 
and the aberrant populations of this subspecies 
from two springs in the Lake Diamond drainage 
basin, but also from G. h. ciicliiUi of Fish Creek 
Springs. Long isolation is therefore indicated, as 
we have already stated. 

A number of springs along both the west and 
east sides of the extensive alkali flat in the north- 
ern part of Newark Valley are shown on the map 
of White Pine County. Nevada ( Ed. Millard & 
Son. Ely. 1930). and on the topographic maps 
cited above. In 1934 we examined springs that 
seemed likely to have maintained native fish. 
Those on the west side, in R. 55 E.. were: near 
Diamond Peak (South Peak), near middle of T. 
20 N. ( Location G7 ) ; at Strawberry Ranch or 
Strawberry, in northern part of T. 21 N.. 18 km. 
south of Simonsen ( where the fish had previously 
died off — see below); and at Moores Ranch, 
about midway between Strawberry and Simonsen. 
as of 1938 (Location G8). On the east, two 
discharging springs are indicated: one. the warm 
spring at Billy Moore's Ranch, close to the T. 
22-23 N,. R. 56-57 E. corner (Location G9 ) ; 
the other, not examined, marked "Sulphur Spr." 
on the 1930 county map. southeast of the center 
of T. 20 N.. R. 56 E. (presumably the "Barrel 
Spring" on the Buck Mounlaiii Quadrangle). 
There are, presumably, a few mountain streamlets 
on the west and possibly on the east side of the 
valley, but the only one that seemed to be pos- 
sibly propitious as a habitat for native fish is 
Cold Creek, which was found to be fishless (p. 
157). 

Characteristics of the springs retaining the na- 
tive chubs in Fish Creek and Newark valleys are 
stated in the accounts of Gila hicolor iic\\(ir(<ciisi.\ 



(pp. 157-158) and G. h. cucliilu (pp. 168-169). 
The springs at Strawberry Ranch, which had be- 
come nearly dry at the time of our visit in 1934, 
v\ere reported by the rancher to have contained 
chubs Luitil they had been completely killed off by 
freezing a lew years previously. 

Pluviai^ Lake Hubbs 

Drainage basin in northwestern White Pine 
County, with the northwestern arm projecting into 
Elko County, eastern Nevada (fig. 7 ). 

Solely on the basis of the crude maps then 
available and local testimony, backed by the rela- 
tionships between enclosed basins and pluvial 
lakes in the general area, we postulated (Hubbs 
and Miller. 1 948b. p. 59) a "lake in Long Valley." 
as probably small and ephemeral, and merely 
mapped (as no. 31) the remnant dry lake. 
Snyder cl ul. ( 1964) correctly mapped a sizable 
Pleistocene lake (no. 52) in the basin, naming it 
Lake Hubbs. Their mapping has been adopted 
by Feth (1961. 1964) and Morrison (1965. 
fig. I ). 

The pointed northern tip of the Lake Hubbs 
drainage basin is inserted between the Lake 
Franklin and Lake Gale sections of the Lake 
Franklin watershed. The eastern edge abuts the 
Lake Gale basin; the southeastern margin, the 
Lake .lake basin; and the western side, the Lake 
Newark and Lake Franklin drainage systems. 

The greatest length of the drainage basin is 90 
km.: that of the somewhat oblong main section, 
roughly 58 km. The greatest width, at a bulge in 
the western border, near the middle of the main 
part, is 32 km. The sharply pointed northeastern 
arm. embracing Long Valley Wash, is about 40 
km. long; its greatest basal width is roughly 18 
km. The form of the lake closely approximated 
that of the drainage basin. The greatest length 
was 51 km.; the greatest width, both in the north- 
ern and .southern sectors, was 14 km.; the least 
width, near middle. 1 2 km. The dimensions of the 
pointed noitheastern arm of the lake were, ap- 
proximately: length. 17 km,; width at mouth. 9 
km.; width near mieldle. 3 km. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



27 



As charted by us, the area of the lake. 490 
sq. km., constituted 29 percent of the area ( L677 
sq. km.) of the drainage basin; roughly agreeing 
with the values ( 205 sq. mi., 32 percent of 635 
sq. mi.) given by Snyder el <//. ( 1964). On the 
basis of our altimeter estimate of 6.266 feet 
(1,910 m.) for the top lake level, repeatedly 
checked against the assigned altitude of 6,352 
feet for the bench mark near the north border of 
T. 20 N., R. 59 E., the depth of the lake is com- 
puted to have been 206 feet ( 63 m. ) above the 
present playa altitude of about 6,060 feet ( 1,847 
m.) indicated by the Ely 1:250,000 map. This 
is somewhat less than the depth of 250 feet listed 
by Snyder et al.. but on a reconnaissance of the 
entire, well terraced eastern shore and northern 
end of the ancient lake, we found no trace of 
higher beachlines. 

Such an accumulation of water is remarkable 
in a valley now almost completely desiccated (fig. 
5 ) . However, the basin is clcsely bordered by a 
mountain rim almost everywhere more than 300 
m.. and in large part more than 600 ni.. higher 
than the 1.847-m. lake bed. On the southwest 
side, only about 30 km. distant to the west, the 
Diamond Mountains rise to an altitude of 3,237 
m. Furthermore, the top lake level probably was 
not attained in the last stages of the Wisconsin 
pluvial: a sample of the fluffy, pure, shell-bear- 
ing marl ( discussed below ) from the top terrace 
deposit yielded a radiocarbon measurement of 
more than 30,000 years B. P. ( Hubbs and Bicn, 
1967. p. 284). Other, lower shoreline features, 
especially terraces and bars near the mouth of the 
northeastern arm of the lake, at an altitude 
mapped as slightly above 6,200 feet ( 1,890 m.), 
therefore at an elevation roughly 45 m. above 
the playa level, definitely have a very late 
Pleistocene appearance. 

Physiographic evidence yields little or no sup- 
port for an hypothesis of any underground contri- 
bution of water from surrounding basins, for the 
playa of Long Valley is slightly higher than the 
flats of lakes Newark and Franklin, respectively 
to the west and north, only about 30 m. lower 
than the playa of Lake Gale, and about 60 m. 




FlouKt 5. Dust rising from the playa on the bed ol 
pluvial Lake Hubhs. in Long Valley. White Pine County, 
Nevada; viewed from eastern shoreline features near 
middle of main part of ancient lake. Photographed by 
senior author September 5. 1965. 

lower than the flat of Lake Jake. Furthermore, 
mountains intervene on all sides. 

The ground waters of Long Valley have been 
described by Eakin ( 1961 ). 

Shoreline and Discharge. 

There is a strong possibility that Lake Hubbs 
once discharged into Lake Newark and thence 
through Huntington Creek and Humboldt River 
into Lake Lahontan. Passes to the Lake Franklin 
and Lake .Fake watersheds are above the 6.500- 
foot (1.981-m.) contours, but the lowest place 
in the sill. Beck Pass, is shown by the Ely 1 : 
250.000 map and the Buck Mountain 15-minute 
Quadrangle to be on the Hubbs-Newark divide, in 
Sec. 10. T. 20 N.. R. 57 E.. where 6,400-foot 
( 1.95 1-m.) contours are separated by about 0.3 
km., above the 6,360-foot (1,939-m.) contours 
on either side. As is indicated below, the highest 
beachline of Lake Hubbs that we could find was 
measured as about 6,266 feet ( 1.910 m.). How- 
ever, the inteidigitation of marl and gravel at the 
bar where this measurement was taken indicates 
fluctuating levels, and the marl may very well 
have been deposited on the exposed shore in 
water of considerably greater depth. That the 
only spilKs) may well have been prior to the last 



28 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 








FifiUKi (i. Intciiicdded lacustrine iiKirl .iiid gr.i\cll\ 
deltaic deposits on eastern shoreline ot pluvial Lake 
Huhbs. in I ony Valle\. White Pine Countx. Nevada^ 
Photographetl h\ senior author Scpieniher 5. I''{i5. 

main pluvial period is suggested by the dating of 
the mail ret'ened to above. The occurrence of 
freshwater moliusks {Gyraiiliis and Pisidiiiin) in 
the marl favors a surface-water connection, but 
the discharge might have been through alluvium 
in the pass, as postulated for Lake Clover ( p. 
30). The hick of fish in Long Valley is not sig- 
nificant evidence for no discharge, because the 
only fish habitat that may have long persisted now 
periodically becomes uninhabitable (see below). 
Limg Valley, known to us previously only 
from maps and local testimony, was explored by 
us I the Hubbscs) on September 4-5, 1965, with 
the great help of the Ely 1:250,000 map and a 
Paulin altimeter reading to 2 feet. The beach 
lines were examined in some detail near the 



middle of the east shore of Lake Hubbs, in T. 20 
and 21 N., R. 59 E. The shoreline features all 
run roughly north-.south, parallel to the lake axis 
and to the contours along the base of the Butte 
Mountains. Going north, we encountered, after 
seeing a slight truncation of the bajada at the 
valley flat, the first definite gravelly beach bars 
in Sec. 29. T. 20 N., R. 59 E., somewhat below 
the mapped 5..iO()-foot ( 1.615-m.) contour. The 
gravel, seemingly stratified and moderately 
rounded, contrasted sharply with the much 
sparser gravel in a road cut nearby. 

About I km. northwest of the bench mark, with 
indicated altitude of 6,352 feet ( 1,936 m.). we 
found, approximately on section line 5-6, in T. 
20 N.. R. 59 E., a recent road-gravel pit obviously 
representing an ancient beach line. Again, gravel 
workings disclo.sed the level of the ancient lake. 
The location was estimated to be 39 3R.2' N. 
lat.. 115 21.9' W. long. Here, for the top of the 
bar. we obtained, by repeated checking of altim- 
eter readings against the bench mark, an estimate 
of the altitude as 6,266 feet (1,910 m.), and 
fecund, close to the eastward, only a slight trace 
of one parallel bar. which registered the same 
altitude. It is c(>nvex toward the playa and ap- 
peared to be about I km. long. There is a weak 
parallel bar immediately west of the pit, and 
traces of other recessional levels farther toward 
the playa. ^ ellow silt deposits were exposed by a 
cut on the east side of the main bar. 

The structure of the bar (fig. 6) had been 
exposed t\ir abinit 6 m. vertically and 40 m. 
horizontally by the gravel excavatit>n. The ma- 
terial was generally coarse, varying from fine 
deltaic gravel, in spots grading into clean sand, 
to coar.se cobble, generally fore.set but in part 
boldly crossbcdded. In one place the gravel 
sharply interdigitated with, and was in part mixed 
with, fluffy silty lacustrine marl, which dissolved 
almost completely in acid. In spots the fine gravel 
was strongly cemented, apparently by marl rather 
than caliche. Fine gravel lenses occurred in the 
layers of marl. Some .streaks rich in nnnute snails 
(Gynnilus sp. ) and some minute clams ( Pisidiuin 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



29 



sp. ) were found in the marl, but no trace of fish 
remains could be found by fracturing the marl. 

Shoreline features were traced in considerable 
detail almost continuously from the southern to 
the northern border of T. 21 N., R. 59 E., over a 
distance of about 10 km. The highest discernible 
traces coincided in level with the bar just dis- 
cussed, and several lower beaches were rec- 
ognized. Across the valley beach features could 
be made out near the base of the hill in T. 20 N., 
R. 57 E., southeast of the low pass. A shoreline 
was observed near the north end of the west side 
of the main body of the ancient lake. Shore fea- 
tures were seen margining each side of the north- 
east arm, again somewhat below the 6,300-foot 
(T,920-m.) contour. 

Close-set bars of relatively recent appearance 
extend in parallel series as gentle curves across 
the mouth of the northeast arm. The most south- 
ern and best defined of these, used as a road grade, 
carries on the Ely 1:250,000 map the elevation 
of 6,205 feet (1,891 m.), and the others were 
indicated by readings of the 2-foot altimeter to be 
at approximately the same elevation. These bars, 
of course, represent the shoreline of the lake when 
it surely was wholly enclosed. 

The Ely and Elko 1:250,000 maps show by 
margined stippling a narrow sandy area 16 km. 
long, in the course of Long Valley Wash, that 
gives the false impression of a flat area. However, 
it is crossed by the 6,300-foot and 6.400-foot 
(1,920-m. and 1.951-m.) contours. The source 
of the sand was apparently not downwash, but 
deposits from dust-devils still seem to be adding to 
the accumulation, for many were seen rising from 
the old lake bed, often to be carried by southerly 
and westerly winds up the trough. These ob- 
viously eolian deposits apparently correspond to 
those at the northeastern end of the Steptoe basin 
(p. 56). 

Remnant Waters and Lack of Fishes. 

Except for flood waters, which at times shal- 
lowly cover the central playa that measures about 
2X5 km. on the Ely 1 : 250,000 map. the only 



surface water on the valley floor is a spring, pro- 
ducing what is known as Long Valley Slough, 
toward the north end of the main body of the 
basin, near the southeast corner of T. 23 N., R. 
58 E. (39 49.6' N. lat., 115" 23.6' W. long.). 
The spring originates in a small patch of tules 
and other vegetation and, when examined on 
September 5, 1965, maintained for about 100 
m. a slight flow containing much CInirci and algae, 
to end in a meadow. A ranch hand at Moorman 
Ranch, who had trapped mustangs at the slough, 
informed us on June 26, 1942, that about July 1 
the water "goes slick" and is then unfit to drink 
and needs to be cleaned out for cattle use. He 
had done so the previous year and was positive 
there were no fish in the spring. Jerome Phalan 
Stratton concurred, and our scrutiny revealed no 
fish. The extreme aridity of the valley is shown 
in a photograph reproduced inside the front cover 
of Eakin's report ( 1961 ). 

There are some mountain springs in the Long 
Valley drainage basin, particularly near the divide 
between this valley and Jakes Valley, but none 
of them can be expected to support fish. One 
of these. North Spring, with a slight discharge, 
was examined in 1965. 

Pluvial Lake Clover 

Drainage basin in southeastern Elko County, 
Nevada (figs. 7, 11). 

This well definable pluvial lake, which has 
been mapped since the time of Russell ( 1885), 
was named and discussed by us ( Hubbs and 
Miller, 1948b, lake no. 26, p. 53), with a review 
of pertinent literature. The name Lake Clover 
has been accepted by Snyder et til. (1964) and 
Feth ( I 964 ) . 

This lake occupied an enclosed basin that is 
largely very flat. The basin is bounded on the 
north by the contiguous drainages of lakes 
Lahontan and Bonneville, on the east by the basin 
of Lake Waring, on the south by the basins of 
lakes Waring and Franklin, and on the west by 
the drainages of lakes Franklin and Lahontan. 



30 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



The maximum, roughly north-south length of 
the western and eastern arms of the dumbbell- 
shaped drainage basin were approximately 59 and 
79 km., respectively, and the corresponding 
lengths of the similarly shaped lake were about 
3S and 46 km.; the east-west dimension of the 
joint basin was about 36 km. 

At its maximum depth of about 42 m. ( sec be- 
low), the lake occupied 930 sq. km., or approxi- 
mately 35 percent of the combined area (2,624 
sq. km. ) of Clover Valley to the west and In- 
dependence Valley to the east. The large extent 
of the lake, with the surface altitude of 740 m.. 
is attributable to the circumstance that more than 
half cif the periphery was closely rimmed by 
mountains rising to altitudes exceeding 2,000 m., 
with several peaks higher than 3,000 m. The 
East Humboldt Range, forming the western rim 
of the basin, is sufficiently high and far north to 
have been extensively glaciated in Pleistocene 
time (Sharp, 1938). Furthermore, Lake Clover 
almost certainly received considerable inflow 
northward from the higher Lake Franklin through 
the alluvium in the low pass between Valley and 
Spruce mountains (p. 42). 

Since the alkali flat in the southern arm of 
Independence Valley has the assigned elevation 
of 5,578 feet ( 1,700 m. ), and the top terrace was 
estimated at about 5,715 feet (1.742 m.), we 
calculate that the maximum depth of pluvial Lake 
Clover approximated 42 m. 

Siioki:line and Dischargu. 

Lake Clover may have attained some subter- 
ranean discharge through alluvium in the pass 
at the north end of the western arm. which was 
only about 30 m. higher than the maxinumi lake 
level, according to altimeter readings and the 
contours on the Wells 1:250,000 map. There 
must once have been a discharge into the head- 
waters of the Humboldt River, however, for the 
valley contains differentiated minnows related to 
and obviously derived from cognates occurring 
in those headwaters, but the spill was almost 
surely during a pluvial period sufficiently remote 



to have allowed the obliteration of the high shore- 
line features by erosion and by deposition. The 
lack of any trace of an outlet channel in the 
gentle, mile-wide pass counters any concept of a 
late-pluvial surface discharge. 

It is quite possible that the outlet level of Lake 
Clover may have approximated the level (about 
5,7 1 5 feet - 1 .742 m. ) of the highest reccignized 
terrace, and that the low sill between the lake bed 
and the watershed of the Humboldt River head- 
waters represents an alluvial aggradation from the 
mountains on either side. For this reason, we 
make no effort to postulate or map a discharge- 
level area of Lake Clover at the lowest altitude 
of the drainage-basin sill. 

Presumably because Lake Clover did not at- 
tain and hold an outlet level in late pluvial time, 
the shoreline features are not very bold. Further- 
more, the western shores of the west arin and of 
the northern part of the east arm are in the lee 
of very high mountains and were, therefore, little 
subject to wave action. In addition, the southern 
shore of each arm was, we assume, protected by 
marsh growth on the gentle slope, in some places, 
however, shoreline features are preserved, as along 
the east slmre. around the tip. and along the east 
shore of the Spruce Mountain peninsula, and on 
the north side of the small island now represented 
as a slight hill marked on the Elko 1:250,000 
map as having an altitude of 5,743 feet ( 1.750 
m.). We encountered, in 1965. on the north 
slope of this knoll, a road-gravel borrow pit where 
a rather thick bed of fine sand and moderately 
rounded gravel indicated an ancient beach, as did 
sand dunes to the northwest and northeast at a 
slightly lower level. Two or three slight rises be- 
tween the borrow pit and the crest of the hill 
(about 0.8 km. distant) seemed to represent 
higher lake levels, but no trace of beach features 
appeared on the top of the knoll, where the gravel 
was angular. Altimeter readings provided an esti- 
mate of 5,710-5.720 feet (about 1,742 m.) for 
the top beachline here. Several beach terraces 
around the north end of the Spruce Mountain 
peninsula, with the uppermost sharpest and with 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



no trace of any higher ones, seemed consistent in 
elevation. The road southward along the east 
side of the peninsula, mapped just below the 
5.700-foot (1.737-m.) contour, closely follows 
sand ridges and some fine gravel ridges indicative 
of old beaches. The gravel pit labelled on the 
Wells map near the north end of Independence 
Valley, at the altitude of nearly 5,700 feet, is 
probably on a near-maximum shoreline. No indi- 
cations of shores substantially higher were seen. 

Remnant Waters and Fish Life. 

Except along a narrow fringe, at the foot of the 
surrounding slopes, the bed of pluvial Lake 
Clover remains as an extremely level alkaline 
flat. A minor part of Clover Valley (the western 
arm of the basin), where enclosed by the 5.600- 
foot ( 1,707-m.) contour (on the Elko map), is 
occupied ephemerally by Snow Water Lake, 
when streams flowing down the draws from the 
southern part of the East Humboldt Range reach 
the bolson. Though the major part of Indepen- 
dence Valley lies within the 5.600-foot contour, 
only "ALKALI FLAT" is mapped there, and no 
water was visible when we were in this valley on 
August 25, 1965, although exceptionally heavy 
rains had recently fallen in this area. The com- 
bined drainages from the many stream courses 
tributary in flood to Independence Valley, in- 
cluding the northern streams of the East Hum- 
boldt Range (see Wells and Elko 1:250.000 
maps), had failed to flood the flat, though con- 
siderable water was standing in the part of the 
valley connecting the two arms. 

There appear to be no permanent streams 
habitable by native fish in the drainage basin. 
One, in Independence Valley, as marked on the 
Wells 1:250,000 map, had no water where we 
crossed it on Highway U.S. 40, August 27, 1965. 
The other stream, marked as permanent, running 
southwestward from near the north end of the 
East Humboldt Range, carries no water as far as 
the valley. There are two large groups of springs, 
each known as Warm Springs, in the basin, one 
cluster on the west side of Clover Valley south 



of the middle and another group on the west side 
of the north arm of Independence Valley. There 
is another rather large spring, on the Wright 
Ranch (formerly the Ralph Ranch) at the south- 
east base of a small hill on the west slope in the 
far-northern part of the north arm of Clover 
Valley. All three of these larger springs contain 
relict fish, discussed below. The three habitats 
are described on pp. 129 and 134-135. The 
waters of the basin were treated, in agreement 
with our findings, by Eakin and Maxey ( 1951b). 

Four glacial tarns are mapped on the Clover 
basin side of the divide between that basin and 
the Humboldt River watershed. They are no 
doubt too high to contain native fish, unless, as a 
remote possibility, trout. 

In the basin we found no trace of Relictiis 
solilariiis, the relict endemic minnow of the 
Franklin-Gale and Waring-Steptoe drainage sys- 
tems that adjoin the Lake Clover basin to the 
southwest, and southeast, respectively. This was 
surprising, since Lake Franklin apparently barely 
missed spilling into Lake Clover (p. 42). How- 
ever, we found evidence that there was a sub- 
terranean discharge, through which fish could not 
have passed. 

Instead, we took in the Clover system samples 
of two endemic subspecies of Rhinichthys osciihis 
(R. ('. olii^oponis and R. o. Icllioponis. pp. 129- 
141) and an endemic subspecies of Gila hicolor 
(G. h. isoluta, pp. 175-180), all of which are re- 
lated to forms inhabiting the extreme upper head- 
waters of the Humboldt River, the main affluent 
of Lake Lahontan. These isolated differentiates, 
as noted above, bespeak a discharge of Lake 
Clover, but at some pluvial period prior to the 
most recent one. 

It is assumed that native fish do not occur in 
the Clover drainage basin at any place other than 
the three just mentioned. The mountain-canyon 
streams are assumed to be devoid of native fish, 
though the possibility of the retention of native 
trout in the streams of the East Humboldt Range 
has not been completely excluded. This possibility 
seems remote, and the explorations and inquiries 



32 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



of trout-specialist Robert J. Behnke and of fishery 
biologists of Nevada have yielded no evidence 
that native trout have occurred in the Lake 
Clover basin. Such canyon streams often fail to 
reach the valley floor, and in the mountains are 
subject to destructively torrential precipitation. 

There is also a possibility, also seemingly re- 
mote, that some cyprinids may have held out in 
some of the other, minor springs along the west 
side of Clover Valley. Mr. Vernon Westwood, 
who had been on Warm Springs Ranch in Clover 
Valley for 1 1 summers, testified in 1965 that the 
only ranches in that valley that contain native fish 
are the three where we found them and the "Der 
Weeks Ranch." which we did not examine and 
concerning which we obtained no information, 
not even its location. 

The valley-bottom waters in Clover Valley are 
almost surely much too impermanent and alka- 
line for fish life. We saw none in the ample but 
obviously temporary waters that had resulted 
from unusually heavy recent rains when, in 1965, 
we examined the area connecting Clover and 
Independence valleys. 

it seems certain that fish are absent thoughout 
the extremely arid Independence Valley, save 
about the cluster of Warm Springs where we took 
the two endemics, Rhinichthys osciiliis Ictlwporiis 
and Gila hicolor isolata. There appear to be no 
permanent surface waters in the Independence 
Valley basin except this cluster and three very 
small groups of springs at the edge of the lake 
bed along the east side of Spruce Mountain Ridge, 
which separates the southern lobes of Clover and 
Independence valleys. On August 25, 1965, 
after failing to find any habitable water around 
the mountainous lobe separating the northern 
arms of the Clover-Independence graben, we be- 
came convinced that no fish exist around the 
southern peninsula. No springs are shown on the 
west shore of this Spruce Mountain lobe. The 
very limited discharge of Spruce Point Spring, 
at the tip of the lobe, was found to be fishless, as 
were the numerous spring sources, mostly mere 
seepages, of Chase Springs. Mound Spring, still 



farther south, was not visited, but a government 
trapper we encountered knew it well and assured 
us that it is a mere seepage, devoid of fish. He 
seemed to be thoroughly familiar with Indepen- 
dence Valley and was certain that the only min- 
nows in the valley are those in Warm Springs 
(mentioned above). 

PLUVIAL LAKES REGARDED AS HAVING 

HAD C ONNECTIONS WITH BOTH 

LAHONTAN AND COLORADO 

SYSTEMS 

( The Lake Railroad Complex ) 

One major and obviously ancient drainage 
basin in central Nevada, that of pluvial Lake 
Railroad, yields physiographic and ichthyological 
evidence of once having had some headwater con- 
nection with the Lahontan drainage system and 
as once having been a part of the Colorado River 
watershed. Two or more now dry and fishless 
adjacent basins, those of pluvial lakes Snyder 
(pp. .^7-38) and Lunar Crater (pp. 36-37), 
furnish doubtful physiographic evidence of having 
been tributary to Lake Railroad. Little Fish 
Lakes (p. 36) definitely did drain into Lake 
Railroad, through Hot Creek Valley, which has 
been held, we think wrongly, to have contained a 
Pleistocene lake (pp. 33-34). 

Pluvial Lake Railroad 

Drainage basin covering several valleys over a 
large area in Nye County, overlapping into south- 
western White Pine County and slightly into 
northeastern Lincoln County, in south-central 
Nevada ( fig. 2 ) . 

This lake, which was probably the largest of 
those between lakes Lahontan and Bonneville and 
which lay in the largest of the drainage basins, 
was named "Quaternary Railroad Lake" by 
Hubbs ( 1*^)4 la, p. 67. fig. 6) and Railroad Lake 
by us (Hubbs and Miller. 1948b. pp. 90-94, 
154, 164). This lake system and its fish fauna 
were discussed by us in some detail, uilh a review 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



33 



of the pertinent literature. The lake has also 
been called "Railroad" by Snyder et al. ( 1964), 
Feth (1964), and others. We now prefer the 
form, "Lake Railroad," to conform with general 
usage for pluvial lakes (p. 3 ) . 

The drainage basin of pluvial Lake Railroad 
is bounded on the northwest and north by the 
basins of ancient lakes Diamond and Newark, 
both of which were once parts of the Lahontan 
system. Along most of the eastern drainage divide 
the tributaries of Lake Railroad abutted those of 
the pluvial White River, a tributary of the Colo- 
rado River; north and south of that watershed the 
Railroad basin is bounded on the east side respec- 
tively by the small basins of Lake Jake and Lake 
Coal, both seemingly dismembered from the 
White River system. Toward the southeast, south, 
and southwest, all separated by rather low divides, 
lie the basins of several rather ill-defined, presum- 
ably shallow, and probably more or less ephemeral 
pluvial lakes, in what we treated (Hubbs and Mil- 
ler, 1948b, pp. 45-46, 149-150, 158-159) as 
the "Area of Sterile Basins," because the region 
seems to be devoid of native fishes and apparently 
was relatively arid even during the very moist 
period. 

The Lake Railroad drainage basin measures 
191 km. in length, from north-northeast to south- 
southwest, and 1 15 km. in maximum width, near 
the middle. The basins of two minor, probably 
once tributary, pluvial lakes, Snyder and Lunar 
Crater (pp. 36-38), form a wedge, from the 
north, which structurally seems to be a southern 
extension of the Newark graben. This wedge 
separates the two main affluents of Lake Rail- 
road. The predecessors of Duckwater, Bull, and 
Currant creeks entered the north tip of Lake 
Railroad. The western affluent, which was by far 
the larger and longer, we named pluvial Russell 
River (Hubbs and Miller, 1948b, p. 92). This 
was obviously a very ancient stream, for its 
course includes two deeply entrenched antecedent 
water gaps. The upper section, the northwestern 
part of the Lake Railroad system, is becoming 
disjunct through the incipient graben depression 



that has formed Little Fish Lakes (see below), in 
Little Fish Lake Valley. Little Fish Lakes 
at high level doubtless discharged through 
a now usually dry structural channel that 
leads into Hot Creek, which follows the deep 
antecedent trench through the Hot Creek Range. 
The hot stream that arises in this trench flows 
at an altitude of about 6,000 feet (1,829 m.), 
between 7,000-foot (2,134-m.) contours sepa- 
rated only 1 .5 km. and between 8.000-foot 
(2,438-m.) contours only 4.5 km. apart (inter- 
preted from the Tonopah 1:250,000 map). The 
flood channel of Hot Creek then debauches onto 
Hot Creek Valley, where it joins a now dry chan- 
nel from the north arm of that valley. After 
coursing through the nearly level Hot Creek 
Valley for 39 km., the pluvial stream entered the 
other water gap, notable but less spectacular, 
through which it passed at an altitude of about 
5.100 feet (1,554 m.), between the 6,000-foot 
(1,829-m.) contours only 4.5 km. apart, in the 
mountain range designated Reveille Range to the 
south and Pancake Range to the north. In high 
flood the pluvial water course is ephemerally re- 
constituted. 

Age-long erosion of Little Fish Lake Valley 
and of the gorge through the Hot Creek Range 
must have led to the aggradation of Hot Creek 
Valley, and sediment from above, supplemented 
by erosion in the Pancake Range, must have pro- 
duced in Railroad Valley the tremendous alluvial 
fan that is marked on the Tonopah 1:250,000 
map by 9 radiating distributaries. This fan, aug- 
mented by another large one, with multiple radial 
distributaries, also debauching onto Railroad 
Valley just to the eastward, has produced, in the 
narrows of that valley, the low saddle that weakly 
divides the basin into northern and southern 
playas. 

These geomorphic considerations have a bear- 
ing on the interpretation of the ancient lakes and 
streams in the area, particularly on the existence 
or age of "Hot Creek Lake" and on the area 
covered by Lake Railroad at its highest stage. 

"Hot Creek Lake." Snyder et al. ( 1964, lake 



34 



CALIIORNIA ACADEMY OF SCIENCES 



MEMOIRS 



nii. 41 ) and Morrison ( 1*^)63. fig. 1 ). mapped a 
Pleistocene lake in Hot C reek Valley and named 
it "Hot Creek Lake." They indicated that it 
covered 88 sq. mi., in a 1,437-sq. mi. basin and 
that it spilled into Lake Railroad. Indeed, the 
area still does so discharge in great floods, but by 
direct surface drainage with no extensive ponding. 
The Tonopah l:2.S().000 map shows the 5.100- 
fool (1,554-m.) contour penetrating through 
most of the gap. whereas the .5.30()-fool ( 1,615- 
m.) contour coincides with the lake margin as 
mapped (except, of course, near the outlet). It 
seems to us that it is unreasonable to conclude that 
any lake, least of all a late Pleistocene one. existed 
here. We saw no trace of lake terraces in Hot 
Creek Valley, nor would we expect any there. 
We interpret the silt deposits in the valley as of 
flood-water, not lacustrine, origin. A very early 
pluvial lake might have existed here. but. if .so. 
we saw no sign of it. and it would seemingly not 
have existed, unless possibly in some much 
earlier hydrographic era. 

Concerning this supposed lake Rush and 
Everett (1966. p. 10) have stated, in apparent 
confirmation of our view: 

.Snyder and others ( 1 ')(i4 ) show ;ni SK 
square-niile Pleistocene lake near 1 wni Springs 
Ranch in Hot Creek Valley that sppjied to 
Railroad Valley. The surtace materials ot this 
area are silt and clay, similar lo those deposited 
in lakes, hut no shore or beach features were 
recognized b\ the writers; theretore the lake 
is not shown on Plate 1 . The log of well 4/ .s 1 - 
13dl (table 1.^) mdicates the presence ot only 
thin beds of lake-deposit type material rather 
than the thick beds usually toiiiul where a 
large and persistent lake occiipieil an area. 

We add that we have often seen sill and clay 
deposits, in places thick, on flat desert areas that 
surely never held a lake. 

Lake Railroad. One or Two? Observations 
of shorelines aking the alluvial slopes well above 
the now dry lake bed of Railroad Valley, near the 
center of T. 8 N.. R. 55 E.. just northwest of 
Locke Ranch, and on the cast side of the valley 



from near Currant to south of Nyala Ranch, 
when reviewed with the Lund 1:250,000 map, 
lead us to place the highest lake shore not far 
below the 5,()0()-foot (1,524-m.l contour. A 
huge longshore, coarse-gravel bar (shown in our 
1948b report, fig. 12), which is just east of the 
road, I 2 km. .south of C urrant. .seems to be, at the 
top. close to the 4.900-foot ( 1.493-m.) contour. 
Since according to observation and local testi- 
mony the saddle between the two dry lakes of 
Railroad Valley is very gentle, and the 5.000- 
foot contours remain well separated here, we think 
it probable that the lake passed through the nar- 
rows, if It did not, at the late high stage, the 
block probably was caused by the conjoining 
aggradation of the two alluvial cones mentioned 
above. We therefore map Lake Railroad as one. 
with an area greater than previously estimated, 
rather than as two. as shown by us in 1948 and 
by Snyder <■/ <//. in 1964 and by Morrison ( 1965, 
fig. 1 ). During late recession stages, at least, the 
lake must, however, have become .separated into 
the two basins. Although the more .southern of 
the two was in Railtxvid rather than Reveille 
Valley, the residual dry lake is labelled Reveille 
Lake on some maps, and Snyder ct cil. applied 
the name Reveille to the lake as a Pleistocene 
entity. If and when separated, the name Lake 
Reveille may be applied to the southern lake. For 
present purposes, however, we treat it as es- 
sentially an integral part of pluvial Lake Railroad. 

On the basis just indicated, we estimate the 
length of the lake ( north-northeast to south-south- 
west), at the highest stage, as 102 km.: the 
greatest width 24 km. in the main area, 10 km. 
in the scnilhcrn expansion. The width in the nar- 
rows was probably only 2 to 5 km. 

The depth of the lake above the altitude indi- 
cated as 4,635 feet ( 1,413 m.) for the present 
playa would have been 340 feet ( 104 m. ) if the 
highest surface altitude was 4,975 feet (1,516 
m. ). We regard this as a good approximation. 
Snyder cl al. { 1964) listed the depth of Lake 
Railroad as 315 feet (96 m.), but gave no value 
for Lake Reveille. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



35 



The area of the lake, as we now chart it, was 
1,394 sq. km., or 13 percent of the 10,874 sq. 
km. of the drainage basin, or 1 1 percent of the 
basin (12,343 sq. km.) if the drainage areas of 
lakes Snyder and Lunar Crater be included. 

In view of the height of the mountains that 
largely border the drainage basin, and slice into 
it in places, this vast accumulation of water is 
plausible, even though the area is transitional be- 
tween the belts of great and only moderate pluvial 
accumulation of surface water. The ridges gen- 
erally rise above the altitude of 2,500 m. and in 
several places exceed 3,000 m. 

The hydrography of most of the basin has been 
outlined by Maxey and Eakin ( 1951 ) and of the 
western part by Rush and Everett ( 1966). 

Remnant Waters and Fish Life in the Lake 
Railroad System, and Probable Past Drain- 
age Connections. 

Our summary (Hubbs and Miller. 1948b, pp. 
90-94) presented in moderate detail an account 
of the dribbling remnant of surface waters in the 
Lake Railroad system, and made it clear that the 
basin is inhabited by only two species of native 
fish, namely a cyprinodontid, Crenichthys neva- 
clae Hubbs ( 1932), which is closely related only 
to Cienichthys bciileyi (Gilbert) of the remnant 
waters of the pluvial White River division of the 
Colorado River system, and a cyprinid (now 
called Gilii bicolor). which is a representative 
of the Lahontan fauna. We further pointed out 
that in only one other pluvial drainage complex, 
that of the Death Valley system in eastern Cali- 
fornia and southwestern Nevada, have the Lake 
Lahontan and Colorado River fish faunas become 
intermingled through vagaries of past hydro- 
graphic connections (Miller, 1946, 1948; Hubbs 
and Miller, 1948b, pp. 77-88, 152-153, 162- 
163, figs. 20-22). We also indicated that present 
topographic relations seem to negate the idea that 
such connections existed during late Pleistocene 
time. The superior topographic information now 
available can be brought to bear on the problem 
of the past hydrographic and fish-faunal connec- 



tions between the Lake Railroad system and other 
drainage basins. 

One possible route that might have been fol- 
lowed in the dispersal of the chub species Gila 
bicolor from its central distributional location in 
the Lahontan system into the Lake Railroad sys- 
tem now becomes evident. The small basin of 
pluvial Lake Snyder (pp. 37-38) straddles a low 
divide between the basins of Lake Newark, a 
former tributary of the Humboldt River still re- 
taining forms of G. bicolor, and the Lake Lunar 
Crater basin, which presumably once, perhaps 
in pre-Sangamon time, led into Railroad Valley 
(pp. 36-37). Very possibly the Lake Snyder 
basin first discharged into Newark Valley and 
then was captured by a former tributary of Rail- 
road Valley through accelerated erosion in that 
direction, that resulted from the circumstance 
that the bed of Lake Railroad is more than 400 m. 
lower than that of Lake Newark. As we have 
previously surmised (Hubbs and Miller, 1948b, 
p. 91 ), this possible dispersal channel may have 
been less remote geologically than the connection 
suggested by the northward-downsloping canyons 
observed in the southern drainage on the Duck- 
water-Newark divide. 

We pointed out in 194S that a past connection 
between Railroad Valley and the Colorado River 
was suggested by the intervening chain of desert 
valleys separated by low divides. The topographic 
information now presented on the Goldfield and 
Caliente 1:250.000 maps and the U.S.G.S. 1: 
500,000 topographic map of Nevada ( 1 965 ) es- 
sentially confirms the postulated route, but sup- 
ports the idea that the connections existed prior 
to the present state of development of the Basin 
and Range geomorphology, for the valleys seem 
separated by divides presumably too high for 
surface-water crossing. The lowest divides out 
of the Railroad Valley depression, lying between 
the 5,600-foot and 5,800-foot (1,707-m. and 
1,768-m.) contours, are southward into the 
cul-de-sac basin of pluvial Lake Kawich and into 
the basin of Lake Penoyer, farther east. From 
Penoyer (Sand Spring) Valley the low pas.ses. 



36 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



between 5,400 and 5,600 feet ( l.(i46 and 1,707 
ni.) lead into Tickaboo (Desert) Valley either 
direetly, or indirectly through the basin of Lake 
Groom in Emigrant Valley. Tickaboo (Desert) 
Valley, with a dry-lake bed altitude oi less than 
1,070 m.. leads via a low-level pass into Pahrana- 
gat Valley, in the course of pluvial White River 
( see the 1 : 500,000 map of Nevada ) . The known 
occurrence of the genus Crenichlhys Hubbs 
(1932) only in the Lake Railroad and pluvial 
White River systems, and the full specific dif- 
ferentiation of the species in the respective basins, 
fits nicely into the hydrographic evidence, as we 
have already pointed out (Hubbs. 1941a, pp. 
66-68. fig. 5; Hubbs and Miller. 1941. p. 2; 
194Xb. pp. 90-93. fig. 24). 

We have already indicated ( Hubbs and Miller, 
1 948b, pp. 90-92 ) that the fish fauna of the Lake 
Railroad system consists of a series of local dif- 
ferentiates of the cyprinid now called Ciihi h'uolor 
(pp. 72. 142) and the entire population of the 
cyprinodont Croiichthys ncvaduc (pp. 227-229). 
The distribution of the various spring populations 
was briefly outlined. More detailed treatment is 
reserved for a future publication. 

Pluvial LiriLE Fish Lakis 

Snyder ct til. (1964). followed by Morrison 
( 1965, fig. 1 ) and by Rush and Everett ( 1966, 
p. 10), showed two small Pleistocene lakes in 
Little Fish Lake Valley, the westernmost arm of 
the Lake Railroad system (fig. 2). Snyder ct iil. 
listed the lakes as no. 5 1 , without assigning a 
name, and estimated their area as 4 sq. mi. ( 10.4 
sq. km. ) in a drainage basin of 467 sq. mi. ( 1,210 
sq. km.). Our measurements are 1 I sq. km. for 
lake area, constituting 1 percent of the area 
(1.131 sq. km.) of the drainage basin, which 
measures 25 ■ 60 km. Snyder cl <;/. correctly 
indicated that these lakes spilled into Hot Creek 
Valley, but only at the maximum stage. The 
Tonopah map shows Fish Lake as permanent but 
little more than 1 km. long, and above and be- 
low it, ephemeral lakes, respectively 2.5 and 1 



km. long. On September 7 of 1934, a year of 
inten.se drought. Fish Lake was an almost com- 
pletely dry alkali flat, but showed signs of having 
overflowed. The two supposed ephemeral lakes 
were .seen to be merely meadow areas. The south- 
ern and the middle lakes apparently together rep- 
resent the larger of the two minute pluvial lakes 
portrayed by Snyder cl nl. and now by us. The 
past and present hydrography of the basin has 
been treated by Rush and Everett ( 1966). 

Ri MNANT Wati:rs and Fish Life. 

The hydrographic and ichthyological evidence 
for this area was briefly treated by us ( Hubbs and 
Miller, 1948b, pp. 91-92). A subspecies of Gila 
hicolor that occurs in springs about the least 
desiccated of the remnants of the Little Fish Lakes 
seems to be closely related if not referable to the 
subspecies occupying Twin Springs in Hot Creek 
Valley and a population in Duckwater Creek (the 
only place where Gila hicolor and Cicnichthys 
lU'vadac .seem to have occurred together). The 
same type has appeared in the outflow of Artesian 
Well 7 on the playa of Lake Railroad, presumably 
by flood connections from either Twin Springs or 
Duckwater Creek, although local testimony, fol- 
lowing common precedent, ascribed the origin of 
the populatiiMi to artesian outflow. 

Pluvial Lake Lunar Crater 

Drainage basin in Sand Spring Valley of Nye 
County, west of Railroad Valley (not to be con- 
fused with Sand Spring or Penoyer Valley in 
Lincoln County); in south-central Nevada (fig. 1). 

The very small pluvial lake that was placed in 
an area carrying question marks on our 1948 
map (because the mapping was inadequate), was 
shown, without being named, as a Pleistocene 
lake on the map by Snyder ct al. (1964), 
which has been repeated by others, also with- 
out assigning a name. It was mentioned, but 
shown only as a playa border, by Rush 
and Everett (19(-)6, p. 10. pi. 1). We pro- 
pose that it be called Lake Lunar Crater, from 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



37 



the name of the most prominent of the several 
conspicuous craters in the area. 

The drainage relations are far from certain. At 
times, the drainage basin may have received a dis- 
charge from pluvial Lake Snyder (see below), 
and Lake Lunar Crater itself may have discharged 
into Lake Railroad. But, on the assumption that 
the drainage was endorheic, as it almost surely 
was at less than maximum lake level, the basin 
was bounded on the cast and south by the Lake 
Railroad watershed and on the west, successively 
from south to north, by the drainage basins of 
lakes Railroad, Snyder (perhaps tributary), and 
Newark; and the pointed north end splits the 
Newark and Railroad tributaries. 

The elongate drainage basin measures 82 km. 
from north to south and, north of middle, 25 km. 
in ma.ximum width (or 29 km. if the basin of 
Lake Snyder be included). As mapped, very 
possibly considerably too small, just beyond the 
margins of the contained lake, the lake mea- 
sured only 2.5 ■ 5.5 km., and covered an area 
of only 1 1 sq. km., less than 1 percent of the area 
of its drainage basin ( 1,321 sq. km. without, or 
1,469 including, the drainage basin of Lake 
Snyder ) . 

Even though the area is now extremely arid, 
and even in pluvial time was presumably rela- 
tively dry, this ratio of lake-to-basin area seems 
incongruously low, especially when we note that 
some high mountains margined the valley. It 
seems plausible to assume that the lake was either 
considerably larger than indicated, or that it dis- 
charged, either on the surface or through alluvium 
or lava, into a tributary to Lake Railroad. The 
discharge may have been temporary or inter- 
mittent, or may have occurred prior to the last 
pluvial. We note on the Tonopah and Lund 
1 : 250,000 maps that the lake basin is separated 
from the Railroad watershed to the south by a low 
and rather broad, flat area in the Pancake Range. 
This flat area is crossed by the 5,800-foot ( 1,768- 
m. ) contour, which just margins the playa that 
is taken to represent the ancient lake. That con- 
tour is mapped as not passing through the low. 



narrow hill ridge (of lava?) to the east, that 
separates the Lunar Crater basin from the Rail- 
road Valley playa. 1,165 feet (355 m. ) lower. 
Local testimony in 1938 confirmed the indica- 
tion that the basin of the playa that we now treat 
as the remnant of Lake Lunar Crater leads south- 
ward across a very low divide into "Mosquito 
Flat" (formerly "Little Round Valley"), which 
in flood drains into Railroad Valley. Whether the 
ancient lake discharged is highly uncertain. 

The hypothesis that a pluvial lake did accumu- 
late in Sand Spring Valley is supported by the 
great extent of the drainage basin, by the defini- 
tively indicated playa, and by the height of the 
surrounding mountain rim, which includes, near 
the north end. Moody Peak. That peak is as- 
signed the altitude of 8,888 feet (2.709 m.) on 
the United States Geological Survey map of the 
State of Nevada 1 : 500.0(30 ( 1965 ) and the height 
of 8,935 feet (2,723 m.) on the Ely 1:250,000 
map. The suggestion on the Tonopah map of a 
cross bar, immediately southwest of the playa 
edge, seems confirmatory. 

Ob.servation of the area by binoculars from 
near the highway (U.S. 6) that crosses the basin 
a few kilometers north of the lake bed ( and north 
of Lunar Crater ) gave no suggestion that any 
permanent waters capable of having maintained 
native fish exist in this basin. 

Pluvial Lake Snyder 

Drainage basin in Nye County, Nevada, north- 
west of Railroad Valley (fig. 2). 

This small pluvial lake was not treated in our 
1 948 report, because we had not visited the area, 
and no topographic map was available. The 
basin was shown, with a contained Pleistocene 
lake, by Snyder ct al. ( 1964) and by Morrison 
( 1965, fig. 1 ), and Rush and Everett ( 1966, p. 
10). but no one has named the lake. We propose 
that it be termed Lake Snyder, as a token recogni- 
tion of the notable contributions that Charles T. 
Snyder of the United States Geological Survey has 
made to the paleohydrography of the Great Basin. 



38 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



The diainagc basin is bounded ou the north- 
west and north by the watershed of Lake Newark: 
on the east and southeast, by that of Lake Lunar 
Crater (just treated); and on the west by the 
Hot Creek Valley division of tiie Lake Raihoad 
drainage basin. 

The basin is irregular in outline and greater in 
north-to-south than in east-to-west dimension, as 
was the lake. The basin is 20 km. long and 13 
km. in greatest width; the lake, as mapped, was 
I 1 km. long and 5 km. wide. 

In the tabulation accompanying their map. 
Synder <■/ <//. ( 1964) listed the area of the lake 
as 10 scj. mi.; that of the basin, as 69 sq. mi.; and 
the depth as "shallow." On the basis of plotting 
the lake boundary along the 6,5()0-foot ( 1,981- 
m.) contour on the Tonopah 1 :250,000 map, as 
Snyder ci al. apparently did, and as seems rea- 
sonable since this contour is shown margining 
what was almost certainly a major longshore bar, 
we estimate the area of the lake, in essential 
agreement, as 24 sq. km., but we estimate the 
area of the drainage basin, by use of said map, as 
somewhat smaller ( 14(S sq. km.). On this basis, 
we compute the area of the lake as 16 rather than 
14 percent of the area of the drainage basin. 

The circumstance that the tributaries came from 
outlying ridges of the high Hot Creek Mountains 
to the west renders plausible the existence of a 
pluvial lake in the basin. More compelling is the 
indication, mentioned above, of a gently curving 
bar extending from the east across most of the 
northern part of the valley flat. 

There is no indication that any other basin dis- 
charged into Lake Snyder, but we note that a 
branch of the main former tributaiy of Lake 
Lunar t rater is shown on the Tonopah and Lund 
1;250,()00 maps as passing within about 1 km. 
of the southern border of the lake. However, we 
may well have mapped the border too low, for the 
contour line ( 6,5()0-foot) used for the lake shore 
is shown on the probable bar, which presumably 
formed under water. It is our provisional and 
questionable hypothesis that the lake at its highest 
level did discharge into Lake Lunar Crater, and 



from there very possibly into Lake Railroad (see 
above ) . 

The possibility that Lake Snyder may have 
discharged northward into the Little Smoky 
Valley arm of the Lake Newark drainage basin, 
rather than into [Railroad Valley, is not excluded 
by the contours on the Toniipah map, becau.se 
the 6,5()0-foot ( 1 ,9S l-m. ) cimtour used to define 
Lake Snyder is not separated by any other con- 
tour from the next 6.500-foot contour to the 
north (the one mislabelled ■"6,700"" on the map). 
In fact, the Lake Snyder basin conjoins the main 
part of Little Smoky Valley and Sand Spring Val- 
ley of Nye County, within a single sublinear de- 
pression. In at least some local u.sage, learned in 
I 93s, the Lake Snyder basin is regarded as part 
of Sand Spring Valley. 

Another rather fascinating possibility, sug- 
gested above (p. 3.^). as an explanation of the 
establishment of the Lahontan species Gila hicolor 
in Railroad Valley, is that the waters in the Lake 
Snyder basin first drained northward into Hum- 
boldt River via the Newark basin, and later were 
captured b_\ the wateis flowing southward, prob- 
ably at some early pluvial perii)(.l. into the much 
lower basin of Lake Railroad. 

Knowing how arid the general region is at 
present, we think it almost certain that the basin 
of Lake Snyder lacks permanent water capable of 
supporting native fish life, and no hint to the 
contrary was given by ouv local infoiniers. 

PLUVIAL LAKES CONTAINING THE 

RELICT DAC E (BETWEEN LAHONTAN 

AND BONNEVILLE SYSTEMS) 

Two enclosed drainage systems, each occupied 
by two pluvial lakes that were connected by stream 
flow, are of central interest in the present study. 
These are the Lake Franklin system, containing 
lakes Franklin and Gale, and the Lake Waring 
system, containing lakes Waring and Steptoe. The 
very minor pluvial Upper Lake Steptoe is inter- 
polated along the course of Steptoe River, the 
main feeder of Lake Steptoe. Lake Antelope, of 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



39 



moderate size, occupied a basin that was pre- 
sumably dismembered from the Lai<e Waring 
basin. 

The Lake Franiciin and Lal<e Waring systems, 
which must have had either some distributary con- 
nection over the intervening divide, or. less 
plausibly, an extremely remote direct connection, 
comprise the entire present native range of the 
relict dace, Reliclus soliUiriiis ( pp. 1 96-207 ) . 

Pluvial Lake Franklin 

Drainage basin in southern Elko County and 
( slightly ) in northern White Pine County, Nevada 
(fig's. 7. ID. 

Pluvial Lake Franklin, one of prime interest, 
both geomorphologically and faunistically. was 
named by Sharp ( I93S ). and was discussed by us 
(Hubbs and Miller. 1948b, pp. 52-5}) with a 
brief review of the pertinent literature. 

This lake occupied a largely very flat basin 
that, so far as is apparent (see below), did not 
quite attain a surface discharge, at least in late 
pluvial time. The northeastern rim of the basin 
adjoins that of pluvial Lake Clover ( pp. 29-3 1 ) . 
The middle of the ea.stern border adjoins the 
basin of Lake Waring (p. 46). The remainder of 
the rim, on the southeast side, is coterminous with 
the north end of the tributary graben of pluvial 
Lake Gale ( p. 45 ) and with the northeastern arm 
of the enclosed basin of Lake Hubbs ( p. 26 ) . On 
the west, the basin is bounded by the great fault- 
block escarpment of the lofty Ruby Mountains, 
the less abrupt western slope of which is drained 
by headwaters of the Humboldt River, the chief 
feeder of Lake Lahontan. Lake Clover was once 
also tributary to the Humboldt, and Lake Hubbs 
may have been, by way of the Lake Newark basin. 
Far to the south the drainage basin of the tributary 
Lake Gale contacts pluvial tributaries to, or basins 
di.sjunct from, the watershed of the Colorado 
River. 

The Lake Franklin drainage basin is weakly 
curved along the western escarpment, is roughly 
pointed at each end. and has a maximum straight- 



line length of 112 km. The longer, western part 
of the basin ( Ruby Valley) is linear, but medially 
the basin is extended eastward, in squarish form, 
to embrace the lower, northern end of Butte 
Valley, which in pluvial time received the dis- 
charge of Lake Gale, through a .short stream that 
we now name pluvial Butte River. The width of 
the central part of the basin is 43 to 57 km. 

The lake itself roughly followed the form of the 
basin. The main, western part. 84 km. long, com- 
prised a south-end expansion about 1 1 km. wide 
and 15 km. long, a connecting strait 5 to 7 km. 
wide and I 1 km. long, and the major body of 
water, which was expanded in roughly triangular 
form eastward. Beyond a constriction only 5 km. 
wide, between projecting points, the lake moder- 
ately expanded into the eastern arm, in Butte 
Valley. Here the impoundment was once thought 
to have been a .separate lake, but it was called 
pluvial Butte Bay by us (Hubbs and Miller, 
1948b, pp. 52-53). This bay, which was hooked 
southward, was 29 km. long, and at its southern 
tip received pluvial Butte River. The greatest 
width of the whole lake was 38 km. 

At its maximum depth of 53 m.. Lake Franklin 
occupied about 1.314 sq. km.. 25 percent of the 
entire drainage basin area ( about 5.27 1 sq. km. ), 
including the hydrographic area of the tributary 
Lake Gale basin (next section), and 39 percent 
of the drainage area (3.338 sq. km.) excluding 
that subbasin. The depth (above present playa 
surface) was estimated by Snyder cr al. ( 1964) 
as 2 1 feet ( 64 m. ). which approximately agrees 
with our interpretation, for the lowest playa eleva- 
tion given on the Elko and Winnemucca 1 : 
250.000 maps is 5,939 feet ( 1,810 m.) and we 
found the highest beach line (see below) to be 
somewhat above the mapped 6,100-foot ( 1.859- 
m.) contour. C. T. Snyder (personal communi- 
cation), by field work in 1969. has placed the 
level as 6.1 13 feet ( 1.863 m.), yielding a water- 
depth estimate of 53 m. 

This tremendous water storage during the 
pluvial periods is attributable to the northern loca- 
tion and to the ereat height of the surroundins 



40 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




5=00' 

WH,'ITE\ I wARiiMG 



Figure 7. Det;iil ot pluvial hydrography of central part of study area ('(" on fig. I ). 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



41 



4.45 




1 15° 00' 

whiteV 

R SYSTEM^ 



^39«I5- 

M°45 

28,29 



Figure 8. Detail of modern hydrography of central part of study area ('C on fig. I ); showing also key geographic 
features, Locations for Gila (G-) and R/iiiiichrhys ( R-), and Collections for /^e/zc/dv (-). 



42 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



nuHintains. Except in a few narrow passes, the 
entire abrupt west rim of the depression lies above 
the 7,000-foot (2.134-ni.) contour, and several 
peaks reach an akitude of 3. ()()() to 3,37S m. This 
range ( Ruby Mountains), along with the contigu- 
ous East Humboldt Range (which contributed to 
the Li\kc Franklin watershed at tiie extreme north 
end), comprised the most extensively glaciated 
area within the Great Basin (Sharp, 1938), be- 
tween the Sierra Nevada and the Wasatch Range. 
Five more or less separated lesser ranges, 
each rising to altitudes above 2.400 m.. and 
several other small and less elevated moun- 
tains, supplied direct inflow from the east, 
and the tributary Lake Gale was almost com- 
pletely flanked by high mountains. 

As is indicated below. Lake Franklin did not 
rise high enough to attain a surface discharge 
through any of the four passes. The divides south- 
ward toward Lake Hubbs. northward then west- 
ward into the Lake Lahontan drainage, and east- 
ward toward Lake Waring, were all definitely too 
high tii have effected an outflow. The pass be- 
tween pluvial Butte Bay and Lake Clover, how- 
ever, was so low that a spillage in that direction 
has been suspected, but apparently was not quite 
attained. In our previous report (Hubbs & Miller. 
l94Xb, p. 52). referring to this pass, we stated that 
our "altimeter readings indicate that at its maxi- 
mum height Lake Franklin here rose to within 
20 feet of the lowest summit, in Wells Fargo 
Canyon" ( the name given on old maps to 
the pass between Valley Mountain and Spruce 
Mountain). The initial reconnaissance, of ,lunc 
27, 1942, which led to this conclusion, was 
repeated by us in more detail on August 
26, 1965. After just passing over the very 
gentle sadtlle southward we were fortunate enough 
to encounter extensive road-gravel works ( at the 
northern tip of Butte Bay as charted on fig. I 1 — 
a point located by compass triangulation ). Gravel 
had obviiHisly been excavated here for fills on 
U. S. Highway 93. Here, well removed horn any 
ancient stream flow, running crosswise of the 
pass, well rounded gravel, rather uniform in size. 
formed a deposit about 5 m. thick. All indications 



are that this beachlinc represents a long stillsland 
of pluvial Lake Franklin. The level seemed to 
correspond with the highest of the beachlines that 
are visible around the entire periphery of ancient 
Butte Bay. including the areas just southward 
and just southwestward. A line of test pits ex- 
tending northward across the summit of the gentle 
saddle (about a mile distant) apparently failed 
to locate a supply of gravel, for none appeared 
in the excavated material, except for fine gravel 
on a low mound about 1 m. high and 10 m. in 
diameter within 100 m. of the divide. This en- 
gendered a seconi.1 thought luitil the same type of 
gravel was found at the surface at higher eleva- 
tions extending up into a draw in Valley Moun- 
tain. This gravel, which markedly increased in 
thickness westward, obviously had been formed 
by stream action on the mountain slope and had 
been deposited largely in an alluvial fan atop the 
divide, with surface sloping downward toward the 
ea.st. Furthermore, the mixture of some coarse 
rounded gravel with much finer gravel and a large 
component of silt did not have the constitution of 
a beach deposit. The mound mentioned above 
and other patches on the flat pass were obviously 
remnants of superficial deposits that had else- 
v\here been eroded away. Many little gullies, 
running in both directions from the divide, follow- 
ing the recent unusually heavy rains, showed that 
during postpluvial time the pass must have been 
subjected to much erosion and presumably to 
considerable lowering in elevation. Therefore, 
the elevation of the pass above the highest Frank- 
lin beachlinc was probably somewhat greater than 
present topography would indicate. 

Closely spaced readings with a Paulin altim- 
eter marked in 2-foot intervals indicated that the 
divide was 44 feet ( 13.4 m. ) higher than the top 
of the gravelly beach deposit ( rather than 20 feet 
as previously estimated). Furthermore, the total 
lack of any outlet channel across the gentle divide 
(the lowest in the margin of the Lake Franklin 
basin ) confirms the assumption that there was no 
pluvial surface spill. We strongly suspect that 
there v^-as a subsurface discharge into the lower 
Lake Clover. Along the canyon leading down to 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



43 



the bed of that lake there are numerous indica- 
tions of higher land levels, suggesting extensive 
stream-channel erosion. Prior speculations re- 
garding the drainage of Lake Franklin, dating 
from before adequate topographic surveys, were 
mentioned by us ( Hubbs and Miller. 1948b. pp. 
52-53). Snyder ct al. i 1964) questionably indi- 
cated a discharge at maximum lake level only. 
The outlet is indicated on Morrison's ( 1965, fig. 
1 ) map. 

The failure of Lake Franklin to overflow is also 
strongly indicated by the lack in the Lake Clover 
basin of Rclictiis solitariiis. the single native fish 
of the Franklin basin, and the lack in the Frank- 
lin basin of the Lahontan derivatives comprising 
the fish fauna of the Clover basin, and of other 
basins now disjunct from the Lahontan system. 

C. T. Snyder's estimate of 6.1 13 feet ( 1,863 
m. ) for the highest level of Lake Franklin (see 
above) is consistent with the height of the top 
terrace all around pluvial Butte Bay and on the 
east side of the southern arm of the main body of 
the lake, particularly on the northwest side of 
Station Butte, where a beachline seemed to be 
visible above the marsh level. Elsewhere along 
the west shore of the ancient lake the pluvial 
beachlines are very poorly developed, presumably 
because this shore lay in the lee of the towering 
Ruby Range. Near the north end of the old lake 
bed, we detected in 1942, from the main road 
crossing to the mouth of Pole Creek ( the northern 
head of Franklin River), seven bars, some well 
defined, on the eastern side. These bars lie at 
altitudes between the 6,000-foot and 6,100-foot 
(1,829-ni. and 1 ,X59-m. ) contours on the Elko 
map (our altimeter readings were consistent with 
these indications). 

Possible Stream Connection Between Basins 
OF Lakes Franklin and Waring. 

Although the basins of pluvial lakes Franklin 
and Waring are separated by a definite saddle en- 
tirely across Goshute Pass, .some means of fish 
transfer across this divide seems to be called for, 
in view of the native occurrence of the distinctive 



endemic minnow. Rclictii.s solitai'ms. solely, and 
with no marked differentiation, in the Franklin- 
Gale and Waring-Steptoe drainage basins. In 
our 1948 monograph (p. 55) we noted that; 

slight drainage channels from the nuuintain to 
the southwest flow over the nearh Hat alkivial 
slope perpendicularly to a definite, though 
small and shallow valle\. which, oddly, 
crosses over the nearly flat divide between 
the two basins. Pluvial streams, possibly large 
and permanent enough to have harbored dace, 
would presumably have wandered back and 
forth across the fan, swinging the drainage 
now toward Butte Valley, now toward Steptoe. 
The little valley crossing the divide looks like a 
stream channel, but if so. it can hardly rep- 
resent a pluvial channel since the divide lies 
far above the highest pluvial lake terraces on 
either side. The trough may have been formed 
by faulting. 

These observations were made near the north end 
of a hill perched on the divide. Another place 
where, much more plausibly, the relict dace may 
have been transferred across the Franklin- 
Waring divide, in either direction, was observed 
by us on August 26, 1965, about 5 km. to the 
northward, where intermittent streams slightly 
entrench the bajada around the south end of 
Spruce Mountain. One channel now leads south- 
ward, then eastward (leftward) toward Nelson 
Creek, which was flooded by Lake Waring, but 
the lay of the alluvial fan here made it .seem al- 
most certain that the channel had flowed at times 
to the right, into Butte Bay of Lake Franklin (see 
Elko map). We strongly suspect that the transfer 
of fish did take place here, but we find no basis 
for favoring a transfer from west ( Lake Franklin 
basin) to east (Lake Waring basin), or a move- 
ment in the opposite direction. RcUcUts now oc- 
curs in spring streamlets about 12 km. distant on 
either side of the divide. 

The occurrence of the highly distinctive relict 
dace in the Franklin and Waring drainage sys- 
tems, and nowhere else, might be cited as evidence 
that these basin complexes were parts of an inte- 
grated river system prior to their separation by the 



44 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



processes that lia\e piiKluced tlie Basin and Range 
physiography. However, since such physiographic 
reorganization, if it occurred, would have been in 
geologically remote time, it would strain plausibil- 
ity to assume that the populations in the respec- 
tive basins would have remained so long without 
any notable differentiation. 

Remnant Waters. 

The ground and surface waters of Ruby Valley 
were treated by Eakin and Maxey ( 1951a). 

Consistent with its location at the foot of high 
mountains, especially the formerly well glaciated 
Ruby Mountains to the west, this basin is. in parts, 
one oi the least arid in the Great Basin. In the 
southern third of the main western part of the 
valley floor there are many ponds and marshes, 
which presently are largely included in the Ruby 
Lake National Wildlife Refuge, wherein hordes of 
waterfowl find protection. These waters, repre- 
senting the ground-water level, very largely cover 
the southern strait mentioned above and the west- 
ern half of the southern bulge of the bed of Ruby 
Lake proper. The lake appears from maps to 
have become disrupted in dry periods, until 
through recent diking and other management 
practices the level on the Wildlife Refuge has 
been more or less maintained. The lake is fed by 
large numbers of small to large springs of clear 
water that issue in file along the obvious fault line 
near the western edge of the valley floor, over a 
distance of about 13 km. Cave Creek, seemingly 
the only permanent stream reaching the old lake 
bed in the southern third of the basin, emerges, 
almost ice-cold. Irom a large limestone cavern on 
the west side. More or less ephemeral playa 
ponds, shown on some maps as "Ruby Lake," 
occur near the north end of the ponds and 
marshes, where the old lake bed opens into a 
wider area. The ephemeral, elongate present 
Franklin Lake in the western part of the valley. 
at about mid-length, is mapped as 13 km. long. 
Occasionally, the bed of this lake is shallowly re- 
filled by the flood-water discharge of several short 
mountain streams from the west and by several 



even smaller and mostly intermittent streams from 
the Ruby Mountains and the East Humboldt 
Range around the north end of the basin. On 
June 28, 1942, Franklin River and Pole Creek 
had an ample flow of clear water well out onto 
the old lake bed, but both tend to dry up there. 

A number of valley springs ( fig. 49) with con- 
siderable discharge tend to maintain Butte Creek, 
on the Lake Franklin (lower, northward-draining) 
section of Butte Valley ( along the course of 
pluvial Butte River), but this streamlet now 
ordinarily disappears before it reaches the exten- 
sive Dry Lake Flat, the floor of ancient Butte Bay. 
The other stream channels debauching onto this 
lake flat are intermittent. The hydrography of 
this entire area has been extensively described and 
mapped in the report by Clancy (1968). The 
general aridity of the sagebrush-covered northern 
Butte Valley, broken by scattered marshy oases, 
is well shown on the cover illustration on the same 
report. Such conditions are rather typical of much 
of the area covered in the present study. 

The various fish habitats in the Lake Franklin 
drainage are dealt with in greater detail in the 
section on Material Examined and Population 
Status, in the account of Relicliis soUtariiis (pp. 
196-207). 

Of the series of glacial tarns near the Franklin- 
Huml~>oldt divide (shown as dots on figs. 7 and 
8), only one, Robinson Lake, the spring-fed 
source of Robinson Creek (in Sec. 23, T. 33 N., 
R. 59 E.). is on the Franklin side of the divide. 
It presumably is too high for native fish. It should 
be checked, however, for native trout, although 
state fishery employees (and we) doubt their oc- 
currence. 

Fish Life. 

The only native fish of the Lake Franklin basin 
is the relict dace. Rclictiis solildiiiis (|ip. 1 96-22(i). 
That this .species is native in the Lake Franklin 
basin is indicated by the following statement by 
King ( 1 S78. p. 504 ) : ". . . . Ruby Lake .... is 
predominantly a carbonate one, but it is of such 
a weak solution that fish are able to live there." 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



45 



The possibility, we think remote, that a second 
species of minnow occurred in Ruby Lal^e was 
suggested by local testimony (p. 20*^)). Rclictiis 
solitariiis very probably occurred, originally, 
throughout the springs and marshes of and im- 
mediately feeding into Ruby Lake, and in the 
springs and their outlets in the northward-draining 
(and also southward-draining) waters of Butte 
Valley (fig. 49). No fish were found in Cave 
Creek in 1934, either within or without the deep 
cave, nor in Franklin River or Pole Creek, when 
they were flowing strongly on June 28, 1942. 
Trout, apparently all introduced, now occur in 
the short streams draining from Ruby Mountains 
toward the old lake bed, and bass and other warm- 
water fishes have been stocked in Ruby Lake, 
where they have very greatly depleted, though they 
apparently have not yet completely eliminated, the 
relict dace. We found no indication that exotic 
fish had been stocked in the springs in the north- 
ward-draining part of Butte Valley. Details re- 
garding the distribution and habitat of Relictiis 
solitariiis arc given in the account of that species 
(pp. 196-208). 

Pluvial Lake Gale 

Drainage basin in north-central White Pine 
County and extreme southern Elko County, oc- 
cupying the major southern part of Butte Valley, 
eastern Nevada (figs. 7. 11, 13). 

This lake, not to be confused with "Butte 
Lake." which turned out to represent a bay of 
Lake Franklin, was discovered and named by us 
(Hubbs and Miller. 1948b. p. 53. lake no. 25). 
The name Lake Gale has been accepted by Snyder 
et al. ( 1964. lake no. 10) and by Feth (1964). 
When the lake was discovered, only the northern 
end was seen, and from the distance its bed was 
erroneously thought to be closed in only about 30 
km. from its north end. Snyder cl al. have shown 
the lake to have been about thrice as long; its 
broadest and deepest section, now represented 
by an extensive playa. is toward the south end. 

The drainage basin of Lake Gale is bounded 
by the watersheds of other pluvial lakes as fol- 



lows: on the north, by Lake Franklin only; on 
the east, very shortly by the Lake Waring division 
and elsewhere by the Lake Steptoe division of the 
Lake Waring watershed; on the south and south- 
west, by tributaries to Lake Jake; and on the west, 
by the drainage basin of Lake Hubbs. 

Both the drainage basin and lake, as is usual 
for minor divisions of the Great Basin, are 
elongated north-south, but both are distinctively 
arched toward the west. The greatest straight-line 
length of the basin is 80 km.; the maximum width 
is 25 km. in the northern part. 29 km. in the 
southern; the minimum width near the middle is 
19 km. Corresponding measurements for the lake 
are 67 km. in length; 8 and 12 km. in greatest 
width, in north and south respectively; 4.5 km. 
in least width, north of the middle. 

At the maximum outlet level, the lake is esti- 
mated to have had an area of 474 sq. km., or 25 
percent of its drainage basin (1,933 sq. km.). 
The greatest depth that the lake attained above 
the present level of the playa in the southern ex- 
pansion of the lake bed. where the Ely 1 : 250,000 
map shows an altitude of 6.161 feet ( 1,878 m.), 
must have been about 35 m., for the outlet level 
was only slightly below the 6,300-foot ( 1,920-m. ) 
contour. The estimate of 80 feet (24 m.) by 
Snyder ct al. { 1 964 ) seems too low. 

So great an accumulation of water is attribut- 
able to the height of the closely margining moun- 
tain ranges, which, except in narrow passes and 
in the narrow northward continuation of Butte 
Valley, consistently rise far above the 7,000-foot 
(2,134-m.) contour, with peaks .shown as high 
as 9.032 feet (2.753 m.) to the west and 10.542 
feet (3,213 m. ) to the east. 

No inflow from any surrounding basin is sug- 
gested, for the drainage divides are definite and 
Lake Gale was higher than any of the surrounding 
lakes except Lake Jake, from which it is separated 
by a pass shown as more than 700 feet (213 m. ) 
higher than Lake Gale. We found evidence, 
which has been accepted, that Lake Gale dis- 
charged northward, over the present very slight 
alluvial saddle in Butte Valley, through pluvial 
Butte River (ca. 12 km. lone) into Butte Bav of 



46 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Lake Franklin. The presence in bdth drainage 
basins of the relict dace. Rcliciiis -.oliuirius. con- 
firms the connection. 

Examination in 1942 of what ue have taken 
to be the northern end of the drainage basin, and 
of the highest of the pluvial lake shorelines (fig. 
49), seemed to leave little dmibt regarding the 
discharge. About 5 km. north and 1 km. west 
of Stratton Ranch on the Elko-White Pine county 
line are two very minor spring sources less than 
1 km. apart that discharge respectively south 
(Twin Springs) and north, on either side of a 
scarcely perceptible divide. About 1 .5 km. farther 
north is the spring source of the northward-flowing 
Odgers Creek. Beginning less than I km. farther 
on. in the gently sloping valley, truncation and 
terracing of the lateral alluvial cones strongly 
suggest the major ancient water course of pluvial 
Butte River. The course extends down the nar- 
row valley between approximated 6.300-foot 
(l.92()-m.) contours as shown on the Elko I: 
2.S0.0()() map. The 6,282-foot (1.9LS-m.) alti- 
tude shown in the northern part of the bed of the 
lake, therefore indicates that here the lake must 
have been very shallow. In the southern basin, 
where the lake-bed altitude of 6.161 feet ( I.S7S 
m.) is shown, the greatest lake depth was pre- 
sumably slightly less than the 150 feet (46 m. ) 
entered by Snyder ct nl. Our approximation is 40 
m. An alkaline lake must have persisted in the 
southern basin after the discharge failed. An 
elongate mountainous island 6 km. long is shown 
in the northern arm. as separated by a narrow 
channel at its north end, but we have no proof 
thai the mountain was not a peninsula. Tenaces 
at the base of a hillock just south of the mouth of 
Snow Creek fix the position of the shoreline there. 



especially along the east and south sides of the 
.south basin. The only waters that are known to or 
would be expected to support fish are the basin 
springs and their short outlets in the far north. 
We fiHind these to comprise three groups, on the 
Wright and Stratton ranches, respectively on the 
north and south sides of the Elko and White Pine 
county line and on the Owens Ranch about 5 km. 
south of the line (fig. 49). Local testimony indi- 
cated that the discharges of these three groups are 
never connected. Twin Springs, about 6 km. north 
of the line, and barely on the Lake Gale side of 
the slight Franklin-Gale divide, were found to 
be nearly dry. Characteristics of the fish-inhabited 
springs are stated in the account of the habitat of 
RcUclus solitaiius (pp. 201-207). 

The hydrography of this area has also been 
described and mapped by Glancy ( 1968). 

Remnant Fish Life. 

The only native fish in the basin is the relict 
dace. Relictiis soliuiiiiis. and it seems to be con- 
fined to the springs just mentioned, in the extreme 
north end oi the trough. The Road Supervisor 
for White Pine County, Jerome Phalan Stratton, 
who had been reared in Butte Valley, told us on 
June 25. 1942. that all mountain springs in the 
area are fishless. An operator on the Stratton 
Ranch on the next day confirmed this testimony, 
and further stated that no fish occur in any of the 
mountain springs or mountain creeks of Butte 
Valley, that all of the valley springs in the basin 
contain "the same kind of minnow with no varia- 
tion." and that carp occur through the valley 
(we took one in the spring creek on the Stratton 
Ranch and noted that planted trout also occurred 
there). 



Remnant Waters. 

The only live waters of consequence in this 
elongate basin .seem to be confined to the north- 
ern arm. Snow and Paris creeks, arising in the 
highest parts of the Cherry C reek Range on the 
east side carry some water, mostly seasonally. 
There are a number of minor nu>untain springs. 



Pluvial Lake Waring 

Drainage basin in southeastern Elko County 
and northern White Pine County. Nevada ( figs. 
II. 13). 

We recounted ( Hubbs and Miller. 1948b. p. 
54) the somewhat invtilved conceptions and 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



47 



nomenclatural history of this major pluvial sump 
lake, for which we proposed the name pluvial 
Lake Waring. This name has been accepted by 
Snyder et al. (1964) and Feth (1964). This 
lake, like most others in the area, persists as an 
extensive alkaline fiat that occasionally holds 
some water. 

The greatly elongated drainage basin of Lake 
Waring, including that of the tributary Lake 
Steptoe, is bounded on the north tip and along 
the northern half of its eastern side by the 
Bonneville watershed; farther south, by the drain- 
age basins of Lake Antelope, which was possibly 
tributary at its highest level, and of Lake Spring; 
at the south end by the basins of pluvial lakes 
Carpenter and Cave, which are physiographically 
part of or are related to the pluvial drainage of 
Colorado River; and on the west, from south to 
north, by the basins of pluvial White River and 
pluvial lakes Jake, Gale, Franklin, and Clover. 
This major north-south fault trough thus inter- 
venes between the Columbia and Colorado river 
systems and bounds on the east the area of minor 
basins treated in this report, with the interpolation 
southward of the parallel trough of Spring Valley. 
The narrow drainage basin of Lake Waring ex- 
tends almost due north from 38 24' to 41 14' 
N. lat.. a distance of 263 km., whereas the greatest 
width was 60 km. and the average width only 
about 30 km. 

The northern part of the basin (Goshute or 
Shafter Valley), which immediately surrounds the 
bed of Lake Waring, is bifurcate southward, where 
it extends on either side of the Dolly Varden 
Mountains. The eastern fork, for which we pro- 
pose the name Antelope Bay, leads southward 
over a weak alluvial divide into the northern part 
of Antelope Valley (as usually mapped; Eakin, 
Maxey, and Robinson labelled the eastern part of 
the main basin "Antelope Valley"). The western 
fork ccinnects through a divide floored with lava 
and alluvium into the tributary Steptoe Valley. 

Pluvial Lake Waring largely filled its basin 
(exclusive of the drainage basins of lakes Steptoe 
and Antelope) and had the same bifurcate form. 



with the eastern fork wider than the western. The 
greatest length of the lake was about 96 km. 
North of the southern expansion the width reached 
18 km. The western fork in a narrow extension 
reached almost to Currie and was probably sepa- 
rated by only 3 or 4 km. from the northern end 
of the tributary Lake Steptoe. The lake narrowly 
transgressed the lower part of the adjacent alluvial 
slopes. One small island rose in the northern part 
of the lake. 

At its maximum depth of about 66 m.. with 
highest water surface altitude ( estimated by 2-foot 
altimeter) of about 5.760 feet (1,756 m.) and 
lowest point indicated on the lake bed of 5,545 
feet ( 1,690 m.). Lake Waring covered 1,314 sq. 
km., which comprised 33 percent of the area 
( 3,949 sq. km. ) of its basin excluding the Steptoe 
basin ( 14 percent of the entire drainage area of 
9,41 1 sq. km., including the Steptoe basin). The 
sharpness of the higher terraces indicates a high 
lake level for prolonged periods. A profuse water 
supply is understandable, since nearly all of the 
closely adjoining mountain crests on either side 
rise to altitudes above 2,100 m. and a consider- 
able number of peaks rear to above 3,000 m. It 
is small wonder that Lake Waring so nearly filled 
its immediate basin, in view of the great volume 
of water that must have poured in through pluvial 
Steptoe River ( including its dilation, pluvial Lake 
Steptoe). Large and prolonged flow is also indi- 
cated by the deep, sharp trench through the lava 
cross-barrier just above Currie, by which Lake 
Steptoe discharged into Lake Waring. As is noted 
below (p. 55), however. Lake Steptoe, accord- 
ing to our calculations, was very much smaller 
than has previously been indicated. Being smaller. 
Lake Steptoe must have lost less water through 
evaporation than it would have at a larger size; 
hence the contribution to the larger and presum- 
ably more permanent Lake Waring was greater. 

There are no indications, either physiographic 
or faunistic. that pluvial Lake Waring discharged. 
As stated below, the lowest passes (through the 
eastern marginal mountains) are at least 40 m. 
hisher than the highest beach lines. Some sub- 



48 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



tenanean outflow through one or more of these 
passes may, through stabiHzation of the lake level, 
have increased the sharpness of the higher ter- 
races. 

It is clear that pluvial Lake Steptoe overflowed 
directly into Lake Waring. In fact. Beckwith 
(1855) stated that Goshute Lake still so dis- 
charged during major floods ( local testimony in 
1938 and 1942, though consistently indicating 
ephemeral filling of the present Goshute Lake, 
was conflicting on this point). Whether Lake 
Antelope of Antelope Valley ever attained surface 
discharge into Antelope Bay of Lake Waring is 
doubtful (p. 60). There are no hints of other 
direct inflows, but a transfer of water and fish 
(Relicliis) by distributary action across the 
alluvial divide between the Lake Franklin and 
Lake Waring basins, in one or both directions, 
seems highly probable (pp. 43 — 1-4). 

Lake Waring Shoreline Data. 

Ancient shorelines, sharply retained along the 
lower slopes of the marginal bajadas of the great 
alkaline flat of Goshute Valley, furnish clear-cut 
evidence of high stands of pluvial Lake Waring. 

The evidence of ancient lake levels is particu- 
larly striking where the rectified shorelines hinged 
on a lava nubbin on the east side of the valley 
4.5 km. east-northeast of Shatter (location shown 
on fig. 12). The hillock was topped by Air Bea- 
con 501, now shown as "Abandoned Beacon" on 
the Elko 1:250,000 map, just above the 5,700- 
foot ( I,737-m.) interpolated contour. The long- 
shore bars (fig. 9) were roughly charted here (by 
C.L.H. ) on .June 23, 1942, by using a hand com- 
pass and a 2-foot altimeter. The bars, largely 
bare or with low herbs, were composed of wave- 
worn gravel and stones. All were elevated a few 
feet and were very regular and parallel. The clay 
soil of the intervening depressions supported deep- 
green sagebrush. As seen from the western side 
of the valley, .shore features extend about halfway 
up toward the level of Silver Zone Pass. 

The western alluvial slope opposite Shafter also 
was seen to be terraced (fis. 10) alon? the road 



<^■si: 



Lava nubbin w 
Air Beacon 501 



'S„ 



^^^y 




^^^^ 


i5-'^.!ir 


.\^ 


=^^on^ 


<\\^®' po^^ 


^0 / 




CP 




, "P 




/ U1 








^ rn 




■ ^ 




> ° 




\- -T] 




' -y 


' 


o 




i. 




a- 


c 


ii 


^ 




--' 


33 






^ 


I> 


' 


Z 


- 


(7) 


• . ^ 


m 


'"■■; n 





Nubbin at 40°52 8' N , I I 4°23 75' W 



Figure 9. Burs on eastern sfiorelinc in northern part 
of Fake Waring, helween Shafter and Silver Zone Pass 
in White Pine Counl\ . Nevada; impinging on a lava nub- 
bin. Hlevations estimated hy use of a 2-foot Paulin 
precision altimeter. From field sketch of June 2.-!. 1^42. 

to Independence Valley, starting with a com- 
puted altitude of 1 .698 m. on lacustrine clay at 
the foot of the alluvial slope, 7.6 km. west of 
Shafter. A particularly sharp terrace 0.5 km. 
farther west and 9 to 15 m. higher, with a deposit 
of wave-rounded gravel, indicated a long low- 
level stillstand. The even slope between estimated 
elevations of 1.739 and 1,751 m. showed traces 
of terraces. Obviously, the lake level was subject 
to much fluctuation. The slope above the defini- 
tive 1,760-m. top terrace was even, without fur- 
ther trace of shoreline features. 

The terraces along the west shore are con- 
spicuous and continuous from near Oasis, close 
to the extreme north end of the valley, almost to 
Currie, including the region south of Dolly Var- 
den Siding on the Nevada Northern Railroad 
where the valley floor gently rises. In general, two 
well defined shorelines are evident toward the top 
level, a very sharp one near the ba.se, and only 
one sharply distinct between. Southwest of Luke 
(north of Flowery Lake) we noticed a fine lake 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



49 




2 5 



Z 



15 
KILOMETERS 



^76 km west of Shatter 

\ Alkoline flat 
698'' 



0.5 



Figure 10. .Shoreline terraces on hajadii on west side of pluvial Lake Waring, west of Shafter, Eiko County, Ne- 
vada. Sketched from quick survey on June 23, 1942. Elevations estimated h\ l^uilin altimeter. 



bar. and the 5.600-foot and 5,700-foot ( 1,707-m. 
and L737-m. ) contours mapped in this region 
(on the Elko map) strongly suggest shoreline 
features. Various other evidences of the ancient 
lake were noted while travelling down the entire 
west side of the valley in 1942. The lake ob- 
viously extended to near the mouth of Phalan 
(not "Phalen") Creek 5 km. north of Currie. 
Whether the intervening distance was occupied 
by a narrow arm of the lake, at its highest level, or 
by the inlet river, is not yet clear. 

The position of the higher lake shores in the 
southern part of pluvial Antelope Bay is indi- 
cated, from notations and sketches made on June 
24, 1942, to be about the same as to the north- 
ward. At the north tip of Kingsley Mountains, 
immediately north of and parallel to the road 
toward Ibapah, Utah, two low terraces parallel 
the road where it skirts a lava nubbin 7.4 km. 
from U. S. Highway 50, and a sharp, even bar, 
made up of beach-worn stones, conjoins the road 
5.3 km. from U. S. Highway 50, for about 2 km. 
Close to the highway a minor terrace runs along 
the road and two more run parallel, just to the 
southwest. The alluvial slope rising toward the 
southwest showed no signs of terracing. About 5 
km. northwest of the highway, near where a corral 
and spring are shown on the Elko map, clear-cut 
terraces were seen on lov\ hills. These beach fea- 
tures appear, by use of this map. to lie between 
the 5,700-foot (1.737-m.) and 5.800-foot 
(1,768-m.) contours. While following the struc- 



tural trough between the beds of lakes Waring 
and Antelope in 1942. it was concluded that 
Antelope Bay pinched out not far from the Elko- 
Whitc Pine county line. This observation, using 
the .same chart, again places the high lake levels 
between the 5,700-foot and 5,800-foot contours. 
The stippled area on the map looks deceivingly 
like a lake bed, but is crossed by the two contours 
just mentioned and obviously portrays sloping, 
sandy, alluvial or eolian, deposits — like those 
shown on the same map to the westward, in the 
Lake Steptoe basin (p. 56). Our mapping of 
Antelope Bay of Lake Waring essentially agrees 
with the highest recognized Pleistocene lake level 
mapped by Harrill ( 1971 ). 

Everywhere around the basin the passes seem to 
be definitely higher than any discernible shoreline 
(estimated at about 1.756 m.). The lowest point 
is probably Silver Zone Pass, shown between the 
5.800-foot ( 1.768-m. ) and 6.000-foot (1.829-m.) 
contours on the Elko map. This pass cuts through 
the Toana Range, on the Waring-Bonneville 
divide. The very flat pass at Cobre. just beyond 
the north end of the basin, on the same watershed 
divide, is shown on the same map to top where 
the 5.900-foot (1.798-m.) contours are very 
slightly separated. The map shows the 6,000- 
foot (1.829-m.) contours barely separated at 
White Horse Pass, through the Goshute Moun- 
tains, also on the same basin divide. All the 
other passes are definitely higher at the south end, 
and the Steptoe-Carpenter divide is shown as 



50 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



LAKE LAHONTAN BASInN-, LAKE BONNEVILLE BASIN 

115.00 II1'45' II4'30' II4°I5' 



IIJ»45- 
4I''I5' 




Fir.UHL II. Detail of pluvial h\Jrograph> o\ northeastern part ol slud\ area ("A' on lig, I). 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



51 



HUMBOLDT R SYSTEM 

IIS-IS' 115-00' 

4ri5' 



DRAINAGE TO GREAT SALT LAKE DESERT 

II4-I5 114*00 




39°45' 



Figure 12. Detail of modern hydrography of northeastern part of study area ("A' on fig. I); showing also key 
geographic features. Locations for Gihi (G-) and Rliinidillixs (R-). and Collections for Relictiis (-). 



52 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



°°'^ 'l franklin ,^J- 



?_::::— 59''45' 







Figure 13. Detail of pkivial hydrography of southeastern part of studv area CB" on fig. I 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



53 



BUTTE VALLEY 




\ MBv^'- 


r" 


^-^'--- A-ii^ > 


1 






-. PRESTdi^-( '{ 




LUND 




( 



Figure 14. Detail of modern hydrography of southeastern part of study area CB' on fig. I ); showing also key 
geographic features and Collections for Rclicrus (-). 



54 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



higher than 7.200 feet ( 2.1^3 in. ). Thus, pliy.sio- 
graphically. the Waring drainage basin is must 
elosely related to that of Lai^e Bonneville, but 
there is no hydrographical or iehthyologieal basis 
for postulating any past smface-water eonneetion 
in that direction. 

RiMN,\NT Waters. 

In view of the tremendous water storage in 
pluvial time and the height of the almost com- 
pletely margining mountains, it is no wonder 
that the drainage basin of Lake Waring is not 
completely desiccated. Most of the surface waters 
are in Steptoe Valley ( these are di.scussed below, 
under the heading of Lake Steptoe). The con- 
nected and possibly once tributary basin ( Ante- 
lope Valley) of pluvial Lake Antelope (pp. 60- 
61 ) is now almost completely dry. 

Cioshute Valley, which was so largely filled 
uith pluvial water, now stands almost com- 
pletely parched. On rare occasions, as during our 
visit in .lime, 1942, much of the great alkaline 
flat is shallowly and briefly covered with water. 
The single, small perennial stream in the basin, 
Phalan C reek, the outlet of Twin Springs on the 
Phalan Ranch, flows eastward to the south end 
of the old lake bed, which it traverses for a con- 
siderable distance northeastward, as NelstMi 
Creek. All other tributaries are short and or in- 
termittent. The hydrography of Cioshute Valley 
(as here interpreted) was treated by Eakin, 
Maxey. and Robinson, 1951 ). 

There are very few springs in the drainage basin 
north of Steptoe Valley proper. Twin Springs, 
just mentioned, emerge from a major bajada, at an 
altitude of 6,200 feet ( 1 ,,S90 m. ) according to the 
Elko map. A group of springs near the junction 
of McDermitt C reek and the ancient outlet chan- 
nel from Lake Steptoe, about .^ to 4.5 km. above 
(south of) Currie. is ditched to that town. The 
only extensive group of springs around the former 
margin of Lake Waring are those on .lohnson 
Ranch on the west side near the north end. about 
7 km. south of Oasis. They lie near the old beach 



line, one on the alhixial slope and others in a 
meadow (as described on p. 20.^ ). According to 
local testimony, the combined waste water may 
run as far as 10 to 16 km., entrenched in the old 
lake bed. The only other spring on or adjacent 
to the lake bed forms what is mapped as "Flowery 
Lake" ( known locally as "Flower Lake"), toward 
the western side of the main body of the ancient 
lake. On June 21, 1942, this "lake" was found to 
be a springfed pool IX ni. long, 0..^ ni. deep, and 
largely choked with vegetation; about 0.4 km. 
north was a chain of small moimd springs forming 
a marshy area about 0.15 km. across (p. 202). 
There are extremely few small springs in the mar- 
gining mountains. Boone Springs in Boone Can- 
yon, tributary to the basin of the eastern arm of 
Lake Waring at an elevation of about 6,300 feet 
( 1,920 m.) irrigated 2 or .^ acres of meadow in 
1942 and was presumably fishless. Other springs 
\sith the same name, in the southein pait of the 
Pequop Mountains (on the west side of the basin), 
were not visited, but like most mountain springs 
are presumably fishless. 

These traces of water in Cioshute Valley are a 
pitiful remnant of the vast pluvial lake that so 
nearly filled the basin. Presumably the high 
Ruby-East Humboldt Range and other mountains 
to the westward have cast a rain shadow on 
Goshute Valley. 

Fish Liie. 

As we have already indicated ( Hubbs and 
Miller. 1 94.Sb. pp. 5.1-55 ), the remnant waters of 
this drainage basin, along with that of lakes Frank- 
lin and Gale, are populated by a single, sharply 
distinct, native species of fish (herein treated 
as R(liclii\ Militdiiiis. pp. 196-226). We believe 
that its original distribution was in springs and 
streams once tributary either to pluvial lakes 
Waring and Steptoe or to pluvial lakes Franklin 
and Ciale, and that it was transferred across an 
mtervening alluvial fan by distributary stream 
flow I p. 43). There is no evidence, however, 
and it ik)w seems highly improbable, that the 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



55 



fish may be a relict dating from some very ancient 
stream system that has become disrupted into the 
two pluvial drainage basins. 

Relictiis solitariiis now seems to be hmited in 
the great sump basin of Goshute Valley to the 
three habitable groups of springs, those just above 
Currie and those on the Phalan and Johnson 
ranches. In addition, as indicated below, it oc- 
cupies many springs and rivulets in the tributary 
Steptoe Valley. No exotic fishes were taken in 
the springs of Goshute Valley, except for rainbow 
and brook trout near C urrie. Several exotics have 
become established in the springs and creeks of 
Steptoe Valley (pp. 236-240). 

Pluvial Lake Steptoe 

Drainage basin in southeastern Elko and 
northeastern White Pine counties, eastern Nevada 
(figs. 7. 11. 13). 

Pleistocene Lake Steptoe. which was named 
"Lower Steptoe Lake" by Clark and Riddell 
(1920. p. 19). and has generally been termed 
Lake Steptoe. is now represented by the ephemeral 
Goshute Lake, in Steptoe Valley above the nar- 
rows just south of C\irrie. This playa sump 
straddles the Elko-White Pine county line. It 
occupies the lowest, northern end of Steptoe 
Valley (in some usages of that name; many 
residents consulted applied the name also to the 
large semi-detached Goshute Valley, the site of 
pluvial Lake Waring, that extends northward 
nearly to Cobre. or at least the southwestern part 
of that valley). Others called the basin of Lake 
Waring "Shafter Valley" and this nomenclature 
appears on some maps. Other maps, including the 
195X edition of the Elko 1 :250,000 map. extend 
the name Steptoe through the intervening narrows, 
but not into Goshute Valley. The 1965 edition 
of the same map and the U. S. G. S. map of the 
State of Nevada ( 1 965 ) carry the label "Steptoe 
Valley" across the whole basin from near Currie 
to near Shafter. and restrict "Goshute Valley" to 
the contiguous, extreme north end of the grabcn. 
between Shafter and Oasis. However. Eakin ct al. 



( 1 967 ) in the state of Nevada water-resources ap- 
praisal of Steptoe Valley, held that this valley 
"extends northward from the southern end of 
White Pine County for about 1 10 miles into the 
southern part of Elko County." and mapped the 
end of the basin at the covergence of the geologic 
contacts, about 10 km. within the hydrographic 
basin of what we term Goshute Valley. Gibbes' 
1873 map (p. 8) labels the main basin 
"Goshute Desert." 

Steptoe and Waring basins are part of a greatly 
elongated trough that extends farther southward in 
Cave Valley, the site of a third pluvial lake. Lake 
Cave, which appears to be structurally related 
most closely to the Colorado River drainage basin. 
Early conceptions regarding Lake Steptoe. in part 
erroneous, and its relation to other pluvial lakes, 
have been recounted by us (Hubbs and Miller, 
1948b. p. 54). 

The drainage basin of Lake Steptoe is bounded 
on the north by that of Lake Waring, to which 
it was tributary; on the east by that of Lake 
Antelope ( a semidisjunct part of Goshute Valley ) 
and that of Lake Spring ( Antelope and Spring 
valleys completely separate the Steptoe and 
Bonneville watersheds ) . On the short southeast- 
ern and southwestern sectors, the drainage divide 
is common with the watersheds of Lake and Cave 
valleys, the sites of Pleistocene lakes Carpenter 
and Cave, respectively. Those lakes were tribu- 
tary to. or disjunct from, the Colorado River 
basin. Beyond the divide on the west side, from 
south to north, the drainage is toward the White 
River ( tributary to Colorado River in pluvial 
time). Jakes Valley (Lake Jake), and Butte 
Valley (lakes Gale and Franklin). 

The Steptoe drainage basin, as we interpret it, 
extends almost due north-south, and is almost 
four times as long ( I 63 km. ) as its greatest width. 
Over most of its length the width is relatively uni- 
form, but varies from 18.5 to 45 km. 

In our present view. Lake Steptoe could not 
have been nearly as large as it was mapped by 
Clark and Riddell (1920), by us (1948b, pp. 
53-55, lake no. 27 ), by Snyder et al. ( 1964, lake 



56 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



11(1. ^)^)), and by others. The two older of these 
maps, which were drawn prior to the topograpiiic 
mapping, exaggerate the outhne much as does the 
1964 map. on which the outhne roughly follows 
the 6,l()()-foot ( l.S.S9-m.) contour as delineated 
on the Ely and Lund 1:25().()0() maps, but with 
fluctuations from 6.(K)() to 6.600 feet. The 1964 
map, however, carries the lake beyond the crest 
of the Steptoe-Waring divide, to put the source 
of the outlet channel 8 km. north-northwest of 
Currie, instead of its actual position 2.5 km. south 
of that town. In so doing, the lake margin when 
redrafted onto the Elko map is seen to intersect 
contours ranging from 6.100 to 6,600 feet ( 1 .859 
to 2.012 m.). The outlet channel according to 
that precise map is between the 5.^M)0-foot 
( 1.798-m.) and 5.8()()-foot ( 1.768-m.) contours. 
When the lake is charted at a little below the 
5.9()0-foot contour, which on other grounds we 
think it approximately reached, its straight-line 
length is indicated to have been only 42 km., in 
contrast to the 110 km. derived from the pub- 
lished maps las measured on the 1964 map). 
The impossibility of even a 6,100-foot lake level 
is obvious when we note that the altitude of Silver 
Zone Pass, between the Waring and Bonneville 
drainage basins, is indicated as being between 
5.800 and 6.000 feet. 

Furthermore. Snyder et iil. attributed to Lake 
Steptoe the wholly implausible depth of .^50 feet. 
Both the outlet channel and the bed of the always 
very shallow, tributary, ephemeral present Lake 
Goshute are mapped as lying between the con- 
tours of 5.800 and 5.900 feet, and we feel sure 
from our field reconnaissance that the difference 
in elevatit)n is considerably less than 100 feet 
(30 ni.). 

Sand deposits exteneling eastward from Goshute 
Lake, then radiating into canyons ( represented by 
a stippled area on the Elko map) may have sug- 
gested to Clark and Riddell and others a lake 
bed. but the area is crossed by the 5.900- and 
6.000-foot contours and extends to the 6.200- 
fool line. Indeed. C lark and Riddell ( 1920, pi. 
2 ) mapped an "Old beach" margining the area. 



and we suppose that this erroneous mapping led 
to the misconception of the lake size. This area 
apparently represents an aggradation of sand, 
transported either by wind or water, or by both. 
A similar but smaller area to the east extends 
along the axis of the northern segment of Antelope 
Valley (p. 49). 

The lake itself, as we now interpret it. mea- 
sured 42 km. in length and 10 km. in greatest 
width. These dimensions are far under those 
stated by Clark and Riddell ( 1920. p. 20). "over 
30 nnlcs [48 km.] long and 11'^ miles [19 km.] 
wide." though they mapped the Pleistocene lake 
bed (on pi. I ) as about 60 miles (97 km.) long, 
as it would be if the lake reached the "Old beach 
line" charted on their plate 2. The area of the 
lake is now computed as 282 sq. km., or 6 percent 
of the area ( 4.646 sq. km. ) of its drainage basin. 
This is much less than the average percentage for 
the basins luidei' tieatment, and only about one- 
third that for the entire Lake Waring watershed, 
including the Steptoe portion. The obvious ex- 
planation for this exception is that Lake Steptoe 
draiiKxl into Lake Waring, which \se now know 
was very much larger and deeper, though it lay 
in a basin that is now much the more arid, and 
presumably also was during the pluvial. Had it 
not been for the discharge. Lake Steptcie would 
no doubt have been nuich Icmger and deeper, for 
it is closely and almost ciimpletcly hemmed in by 
moLintains mostly rising to altitudes higher than 
2.500 m.. and attaining elevations higher than 
3.000 m. in several peaks, in both the Schell 
Creek and Egan ranges. 

Because Lake Steptoe was much shorter than 
it has commonly licen thought to have been, the 
lengthwise pluvial Steptoe River, as we have 
called it ( Hubbs and Miller. 1948b. p. 54). was 
correspondingly longer than pieviously indicated. 
At the height of the late pluvial it was no doubt a 
major stream. 

It has been observed, by us and others, that 
shoreline remnants are relatively inconspicuous in 
Steptoe Valley. The probable reason is that the 
lake, because of the altitude of its discharge chan- 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



57 



iiel, did not rise far onto the gentle slopes of the 
valley floor, and the marginal shoals were in 
all probability so well protected against wave 
action by a heavy growth of marsh vegetation 
that sharp terraces or bars were not formed. 
Within a single depression, for example that of 
Lake LeConte in the Salton Sea basin of south- 
eastern California, we have found that the bold- 
ness of shoreline features on steep slopes contrasts 
very sharply with their almost complete absence 
on very slight slopes. 

Since all passes in the rim of the basin of Lake 
Steptoe, except the outlet into Goshute Valley, 
are indicated by the Elko. Ely. and Lund 1 : 
250,000 maps to be more than 200 m. higher than 
Lake Steptoe as now delineated, and since there 
are no indications that any of the lakes in ad- 
joining basins rose high enough to crest the 
divide ( indeed all of them drained or would have 
drained elsewhere), there was no inflow, at least 
of surface water, from any of the surrounding 
depressed basins. In this conclusion, we disre- 
gard pluvial Upper Lake Steptoe (p. 59). which 
merely lay in the course of pluvial Steptoe River, 
and the lake ( no. 27 ) that Snyder ci al. ( 1 964 ) 
indicated as of Pleistocene occurrence in Egan 
Creek Valley, but which we think could not have 
existed there, at least in kite Pleistocene time 
(p. 58). 

There is clear evidence of the discharge of 
pluvial Lake Steptoe, between the north end of 
the present Steptoe sump (Goshute Lake) and 
Currie (a straight-line distance of about 7 km.). 
Examination of this area on August 24 and 25, 
1938, and also in 1934 and 1942. disclosed a 
definite stream channel, steeply entrenched in bed- 
rock about 2 to 3 km. south of (above) Currie. 
that served as the course of the Nevada Northern 
Railroad ( Hubbs and Miller. 1948b. p. 54). It 
appears that the large alluvial cone formed by 
McDermitt Creek and smaller draws, from the 
west, had in past time, probably early pluvial, 
blocked a former outlet channel, so as to force 
the stream onto bedrock, which was then en- 
trenched. Rejuvenated erosion extended up Mc- 



Dermitt Creek, to leave the rather strong lateral 
terraces that are visible there. The lateral alluvial 
cones were truncated by the outlet of Lake Step- 
toe. A now dry river channel that is obvious from 
Currie to the mouth of Phalan Creek ( about 6 
km. ) probably formed after Lake Waring dropped 
below its highest level to lay bare the bottom of 
the narrow north-end tongue of the basin. The 
Elko map shows the 5,800-foot ( 1,768-m.) con- 
tour extending as a V into the outlet channel to 
about 2.5 km. south of Currie. but the inter- 
polated 5,900-foot (1.798-m.) contours arc 
separated throughout. 

Rkmnani WaTJiRS. 

Lake Steptoe is represented by the remnant 
sump. Goshute Lake; by an axial, more or less 
continuous tributary thereto; by a number of other 
small, mountain and valley streamlets; and by 
many springs. 

As already mentioned, Goshute Lake is an 
ephemeral body of water at the extreme north 
end of the basin of pluvial Lake Steptoe. It is 
shown on the Elko map as narrow and 13.5 km. 
long, with a marshy extension southward, fed 
by Duck Creek and lateral streamlets. 

The main axial stream in this valley, the rem- 
nant of pluvial Steptoe River, usually carries some 
water, even in dry seasons, despite the use of most 
of the available water by the mills at McGill, as 
Clark and Riddell (1920) noted long ago in 
describing the streams and springs of the valley. 
About a century ago explorers reported that this 
stream required bridging for wagons to pass, 
though it was said to go dry later in the season. 
The stream is labelled Duck Creek on most maps, 
though locally wc found it usually called "Steptoe 
Slough." On the Ely 1:250,000 map ( 1959) it 
is labelled Duck Creek on its lower half, then 
Murray Creek just below Ely. It is labelled Step- 
toe Creek just above that town, and along the 
watercourse that debauches from the Schell Creek 
Range and flows westward to join the axial stream 
course at a right angle. Then, farther south, above 
a marsh just south of the abrupt bend in "Steptoe 



58 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Creek," the axial stream is again labelled Murray 
Creek on the map. However, the Ely I 5-minute 
Quadrangle labels the axial stream Willow Creek 
below the point where Willow Creek proper joins 
the axial course at a right angle. All the axial 
courses, however named, wc regard as remnants 
of pluvial Steptoe River. Except during torrential 
precipitation the streams on the valley floor are 
mostly small and slow. There are also a consider- 
able number of more or less permanent streams. 
in the canyons of the adjacent mountains, that are 
devoid of native fish. 

In addition io numerous mountain springs, a 
number of valley springs extend from 92 km. 
north to 10 km. southeast of Ely: these locations 
range from 1 cS km. south of the north end of the 
basin to 4S km. north of the south end. From 
north to soiUh the main valley springwaters are: 

Springs on the HIko C oiinl>' sale o! the HIko-VVhitc 
Pine count)' line. 

•Springs on the C;irdano k;nKh. leecling the marshes 
just siHith of (loshute lake, ,ind minor springs in the 
s.ime general regii>n. all on the west side. 

Alter a long gap (nearly 4(1 km.), NUmte Neva Hot 
Springs ("Melvin Hot .Springs" ol Clark and Riddell, 
U)2()), with cooler outflow below. 

Springs on the Campbell or Steptoe Ranch, and, just 
hevoiid, Cirass Springs, with many minor springs in file, 
issuing along a probable fault line in what Clark and 
kiddell (1^20) and Eakin <7 al. ( I ')(,7 1 called the 
"Campbell Embayment." 

Springs on Dairy Ranch neur McCnll. the tirst I rom 
the north on the east side of the valle\ 

Spring water at Georgetown Ranch, near l:ly. 

Finally, springs on the CCC (Consolidated Copper 
Company), now C-B Ranch, in mid-valle\, where 
Sleploe Creek turns .ihruptly northward on its .illuvial 
Ian. 

Farther south, in the part of the basin that discharged 
through Upper Steptoe Fake, isolateil mountain springs 
aiKl a cluster of cool outflows in Willow Creek Basin — 
.ill seemingly devoid of native tish (p. dO). 

All of these spring waters are discussed in more 
detail in the account of the habitat of Reliciii.\ 
solihiriiis (pp. 204-206). The hydrography of 
Steptoe Valley has been treated by Eakin ci al. 
(1967), 



Fish Life, 

The only native fish in this drainage, as also 
in the watersheds of lakes Waring. Gale, and 
Franklin, is the relict dace. Relictiis solitariiis. 
It occurs, usually in great abundance, almost en- 
tirely in the warmish ( neither hot nor cold) valley 
springs and in the springfed valley creeks and 
sloughs. The mountain springs and creeks seem 
to be avoided by native fishes. The only seemingly 
suitable spring waters where we failed to find this 
fish are those on the Murphy, formerly Dolan, 
Ranch on the Elko C cuinty side of the Elko- 
White Pine county line. Here, and in marsh be- 
low, it probably once abounded, according to 
local testimony (p. 206), but it is now replaced 
by the obviously introduced Utah chub. Gila 
atraria (p. 2.i 1 ). Other introduced fish found in 
the basin are rainbow and brook trout, carp, gold- 
fish, carp goldfish hybrids, and Sacramento 
perch (pp. 2.^5-240), 

The Supposed Lake in Egan Basin, 

A second lake ( no. 27 ) tributary to Steptoe 
Valley was indicated by Snyder ct al. (1964), 
followed by Morrison (1965, fig. I), to have 
existed in the nearly enclosed Egan Basin, That 
basin lies just west of the mid-length of the Lake 
Stepttie bed. between the narrow northern end 
of the Egan Range and the southern end of the 
Cherry Creek Mountains, with a relatively low 
pass extending southward to the southern basin 
of Butte Valley, that is. toward what was the 
deepest part of pluvial Lake Gale. The lake has 
not been named, but was listed by Snyder <7 cil. as 
having had an area of 10 .square miles in a drain- 
age basin of 62 square miles. However, the topo- 
graphic features, as shown on the Ely 1 : 250,000 
map, provide little basis for the hypothesis that a 
pluvial lake existed in this basin. The lake as 
mapped extended to just within (below) the 
6,800-foot (2,07.^-m.) contour and covered the 
6.600-foot (2.()12-m.) and 6.400-foot ( 1 .95 I -m.) 
contours, both of which passed without closing 
through the very narrow and steep outlet canyon 



VOL. Vll HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



59 



of Egan Creek, intti which canyon the 6,200-foot 
( 1,890-m. ) contour inserts from Steptoe Valley. 
Any ponding of water in this basin that could 
have occurred would necessarily have been prior 
to the deep erosion of the canyon, presumably 
during an erosional epoch that preceded the late 
pluvial period with which we are concerned. We 
question whether any lake ever existed here, but 
have made no pertinent field studies. The hydrol- 
ogy of Egan Creek was briefly described by Clark 
and Riddell (1920, p. 34). The canyon was 
traversed by the old Overland Mail Route (p. 
235). 

Pluvial Upper Lake Steptoe 

Drainage basin comprising the southernmost 
fifth of Steptoe Valley, in south-central White 
Pine County, eastern Nevada (fig. 13 ). 

Clark and Riddell ( 1 920, p. 20 ) briefly indi- 
cated that a small and shallow "ancient lake" ac- 
cumulated in this area, and they assigned to it a 
definitive name, "Upper Steptoe Lake," to dis- 
tinguish it from the main "ancient lake" of the 
basin, which they named "Lower Steptoe Lake." 
We ( 1948b. pp. 54-55. map 1 ) mapped but did 
not name or discuss the southern "lake." Snyder 
('/ ill. (1964) mapped the lake (their no. 100) 
as Pleistocene, but indicated it as unnamed (de- 
spite Clark and Riddell's proposal). Snyder ct 
III. designated the lake area as 10 square miles 
(26 sq. km.) and the drainage-basin area as 358 
square miles (927 sq. km.), and stated that it 
drained into Lake Steptoe ( an obvious conclu- 
sion). Our estimates are 13 sq. km. for the lake 
area (not precise), less than 2 percent of the 
area. 816 sq. km., of the drainage basin. 

We provisionally and rather doubtfully include 
the lake as of Pleistocene age. and name it. in 
slight emendation. Upper Lake Steptoe. reserving 
the unmodified name Lake Steptoe for the larger 
and more definite impoundment to the north. 

As Clark and Riddell indicated, it appears that 
the "ancient lake" — and this would seem to be 
true, even though it was merely a pond, or even 



a marsh — accumulated above the narrows in the 
valley, where the axial drainage was slightly 
blocked by alluvial aggradation from the upper 
segment of Steptoe Creek on the east side and 
Sawmill Creek on the west side. These authors 
logically added that "the lake was apparently 
drained by the dissecting of this alluvial divide." 
We fail to see how a lake of considerable depth 
and size could have accumulated here, at least in 
late pluvial time. However, the evidence of a 
cross-basin barrier suggests that the extensive 
meadows may represent the silted-in remnant of 
an ancient lake, of shallow depth and about 10 
km. long by about 3 km. in greatest width. The 
maximum primary dimensions of the drainage 
basin are 23 ■ 48 km. The lake and basin arc 
shown on the Ely 1:250.000 map. the Ely 15- 
minute Quadrangle, and the Comins Lake 7.5- 
minute Quadrangle. 

Remnant Waters. 

The ancient lake is represented, we think 
largely by reason of earthen impoundments, by a 
variable body of water extending about I km. 
regularly, and about 2 km. rarely, southward from 
near the abrupt northward turn of Steptoe Creek, 
as is shown on Clark and Riddell's map ( 1920, 
pi. 2) and on the quadrangles mentioned above. 
According to local information, the southern, in- 
termittent area filled in 1969. as a result of un- 
usually heavy snow and rain, for the first time 
for 35 years. The highway embankment was 
about 3 m. higher than the small, cutoff northern 
piece of the main pond. No shoreline indications 
of a pluvial lake were seen. 

The modern lake was named Cummings Lake 
by Clark and Riddell (1920). who also dealt 
with Cummings Ranch and Cummings Spring, 
north of Ely in the same valley. The Ely 1 5- 
minute Quadrangle ( 1 958 ) uses the name Comins 
Lake, as does the Comins Lake 7.5-minute 
Quadrangle; the U.S.G.S. State of Nevada map. 
I : 500.000 ( 1965) gives the name Comins Mea- 
dow to the area just south of the modern lake 
(unnamed). Local testimony seemed to indicate 



60 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



the spelling of Cummins or Cumin Lake. We 
assume that Comins is the eonect speUing. for 
we note that the History of Nevada, edited by 
Myron Angel (1881). provided a biographical 
sketch and figure of Hon. Henry A. Comins. who 
settled in Nevada in 1863, removed to White 
Pine [County] in 1869. and engaged in lumber- 
ing, farming, and mining. 

The two streams of the drainage basin, both in 
part permanent, arise in the Egan Range. The 
main one. Willow Creek, which rises in the pro- 
fusely spring-fed, relatively broad and flat Willow 
Creek Basin, on the eastern slope of the range, 
near the north border of Sec. ,^5. T. 14 N., R. 
63 E.. irrigates Willow Ranch, and contributes 
some flow to the valley axis, but becomes inter- 
mittent above Comins Lake. Farther south. Wil- 
low Creek is paralleled in its eastward flow by 
Williams Creek, which is intermittent in the valley. 

Fish Life. 

No trace of native fish was found in the drain- 
age basin by a reconnaissance on August 16-17. 
1969, although Relictiis abounds in Fish Pond 
Springs immediately below, in the bend of Steptoe 
Creek {Collections 28 and 29). Spring-fed Wil- 
low Creek appeared from an inspection of the 
maps to be a propitious site for Rcllclus. but the 
springs and their outflow were found to be in- 
habited only by introduced trout. The inspection 
disclosed no apparently suitable habitat for the 
relict dace, which ordinarily avoids cold springs 
and nnumtain streams. 

Comins Lake was found, unexpectedly, to 
swarm with the Utah chub, GiUi alnirin. obviously 
as a result of a recent, previously unreported in- 
trodLiction (pp. 231-232). 

Local testimony indicated that trout and bass 
(presumably Microptenis salmoiclcs) were stocked 
in Comins Lake in the spring of 1969. 

Pluvial Lakf. Antelope 

Drainage basin in northeastern corner of White 
Pine County, eastern Nevada (figs. 11. 13 ). 



This minor pluvial lake was named by Jones 
( 1940) and the confused history of its drainage 
relations were recounted by us (Hubbs and Mil- 
ler, 1948b, pp. 55-56). The situation is now 
considerably clarified by use of the Ely 1 :25().()()() 
map. The lake undoubtedly occupied a .separate 
fault-block depression, though it lies in the trough, 
long known throughout as Antelope Valley, that 
continues into or even beyond the southeastern 
fork of the Lake Waring basin. The Lake Ante- 
lope basin has been mapped as Tippett Valley by 
Harrill ( 1971 ) and by Rush et al. ( 1971 ). The 
saddle that splits the drainage that in pluvial time 
flowed northward into Antelope Bay of Lake 
Waring (p. 49 ) and southward into Lake Ante- 
lope, is attributable not only to the close approxi- 
mation of the slopes of the Goshute Mountains 
and the Antelope Range, but also to the develop- 
ment of a major alluvial fan from the main stream 
course in the Antelope Range. The fan is 
delineated on the Ely map by ditches radiating 
downward to supply both watersheds. 

The pluvial-lake drainage basins surrounding 
the Lake Antelope basin are Waring to the north, 
Bonneville to the ea.st, and Spring to the south 
and west ( just missing near-contact with any head- 
waters to Lake Steptoe). 

The roughly oblong drainage basin measures 
47 km. long by 27 km. in greatest width. The 
corresponding dimensions of the centrally located 
lake were probably 23 and 7 km. The lake bed 
is deepest near the middle, u here a pond accumu- 
lates (Elko 1; 250,000 map). 

As we have sketched it, largely on the basis of 
field reconnaissance (C.L.H.) of June 24. 1942, 
and on an examination of the Pleistocene lake 
map of Snyder ci al. ( 1964), Lake Antelope 
covered at its highest stage 125 sq. km., or 14 
percent of the area ( 870 sq. km. ) of its drainage 
basin. Its greatest depth was estimated by those 
authors to have been 75 feet (23 m.). Its outline 
seems to have run slightly above the 5,700-fool 
(1,737-m.) interpolated contour on the Ely I: 
250.000 map. The rough correspondence with 
the maximum height of Lake Waring, in the same 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



61 



structural trough, elicits the thought that the two 
pluvial lakes may have had an underground con- 
nection through an alluvial block, or possibly that 
the waters were confluent and that the alluvial 
aggradation was postpluvial. 

Oddly, the rather flat mountain passes between 
the pluvial basins of Lake Antelope and Lake 
Waring, between Lake Antelope and Lake Bon- 
neville, and between Lake Waring and Lake Bon- 
neville, are all mapped as cresting between the 
5,800-foot (1,768-m.) and 6,U00-foot (1,829-m.) 
contours (as drawn on the Elko 1 : 250,000 map). 
When these passes were all checked in 1942, no 
indication was found of any surface-water pluvial 
discharge. The approximate agreement in pass 
altitudes renders almost hopeless any attempt to 
reconstruct any possible remote-pluvial drainage 
pattern. 

The 1942 field reconnaissance, taken on the 
old Overland Mail Route, indicated that definite 
shorelines persist around the outlines of pluvial 
Lake Antelope. Tlie field record provided ma- 
terial for the following revised description of shore 
features, in order of observation: 

0.0 km.: Approaching on the road from 
Deep Creek (Ibapah), arrived between two 
beach lines about 6 m. apart in elevation and 
about 0.15 km. on either side of tine road 
[these beaches presumabh' lie on the gently 
curved ridge, followed by the road, shown 
below the 5.700-foot (1.7.^7-m.) contour on 
the Elko map|. Below, on the lake flat, are 
rather large dunes. 

2.9 km.: On definite terrace 43 m. below 
Antelope-Bonneville divide (by altimeter, not 
rechecked ) ; some standing water on the lake 
sump. 

3.7 km.: Bench extended from mountain 
onto flat as a definite though somewhat com- 
plex bar, which ran 2.7 km. before merging 
with bajada from Antelope Range. This bar 
was separated from mountains along its course 
by a lower flat, in places as wide as 1.5 km.; 
from which flat, drainage had broken through 
the bar in two places. Water-worn stones were 
found on the bar where it was followed by 
the road. 

7.2 km.: Junction with road down Ante- 



lope Valley from north; beach lines observed 
just to north. 

1 2.4 km. : Arrived at Tippett ( 49 m, below 
Antelope-Bonneville divide by altimeter; not 
rechecked ). 

13.7 km.: About 7 m. below distinct wave- 
cut terrace, truncating alluvial cones. 

18.2 km.: Near end of sand dunes. 

These observations seem consistent with the 
mapping of the lake and its drainage basin by 
Snyder et al. (1964) and by Harrill (1971). 
Some features shown on the Ely 1 : 250,000 map 
presumably represent bars and dunes. 

Remnant Waters and Apparent Lack of 
FrsH. 

Throughout Antelope Valley, both north and 
south of the divide that separated the drainages 
into lakes Waring and Antelope, we saw no 
springs that could possibly be expected to harbor 
fi.sh. The region about Tippett is irrigated by 
water piped and ditched from a warm spring in 
the Antelope Range, but we saw no fish in the 
ditches. This was reputed to be the only per- 
manent water .source in the basin. 

Local informers, familiar with native creek 
fishes in Deep Creek ( in the drainage basin of 
Lake Bonneville, just beyond the intervening 
divide), stated that no fish occur in the Tippett 
area. In all probability there are none, and none 
may have occurred even when the pluvial lake 
filled the lower part of the basin during late 
Pleistocene time. At an earlier pluvial period 
lakes Antelope and Waring were possibly con- 
nected, and fish then may have occurred in the 
Lake Antelope basin. 

PLUVIAL LAKE SPRING (SEPARATED BY 

LOW DIVIDES FROM BOTH BONNEVILLE 

AND COLORADO SYSTEMS) 

One trenchlike depres.sed basin that included 
a pluvial lake, namely that of Lake Spring, may 
represent an ancient disruption of either the Bon- 
neville basin or the pluvial Carpenter River, and 



62 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



hence also ihe W liite River division, nf the C olo- 
rado River drainage system. Untortunately. it 
seems not to have retained any native fish fauna, 
for any such remnant would almost surely have 
provided a clue to the past hydrographic connec- 
tions. 

Drainage basin extending through nearly all 
of the north-south extent of eastern White Pine 
County and into northeastern Lincoln County, in 
far-eastern Nevada ( figs. II. 13). 

This lake was named by Jones (1940) and 
was briefly discussed by us ( 1948b. pp. 56-57). 
Its floor is a long, flat valley, with a major, oc- 
casionally dry. sump locally called White Lake, 
north of the middle of the basin. At the south end 
is the narrowly connected Baking Powder Flat, 
containing "The Seep." 

The basin of Lake Spring is bounded on the 
north end by the drainage of Lake Steptoe: on the 
east side, by the watersheds of lakes Antelope and 
Bonneville; at the extreme south cm\ and part 
way up the west side, by the tributaries of Lake 
Carpenter; throughout the rest of the west side 
by the drainage basin of Lake Steptoe. The basin 
of Lake Spring thus intervenes between the major 
hydrographic systems of Lake Waring. Lake Bon- 
neville, and the Colorado River. 

This greatly elongate enclosed fault basin ex- 
tends almost due north-south. It resembles and 
parallels the Lake Steptoe basin, but starts and 
ends farther south. Its greatest length is 180 km. 
and its greatest width, north of the middle, is 42 
km. A stretch as narrow as 21 km. lies south of 
the middle. The lake was 96 km. long and its 
maximum width was only 17 km. in the northern 
and 1,^ km. in the southern part. Between these 
two parts there appears to have been, at the 
highest lake level, a strait narrowing to about 
3 km. 

Lake Spring at its deepest filling was a large 
body of water, covering 978 sq. km., as we mea- 
sure it. C. T. Snyder and associates, accepting a 
less elevated (and more certain) highest shore- 
line (see below) computed the lake area as 332 
square miles — 860 sq. km. ( Snyder (7 <//.. 1964) 



or 335 square miles = 868 sq. km. (Snyder and 
Langbein. 1962, p. 2385). The corresponding 
estimates for the area of the drainage basin are 
4,337 ,sq. km., 1,641 square miles - 4,253 sq. 
km., and 1.630 squale miles = 4.222 sq. km. 
According to these figures, the lake area com- 
prised 23, 20, or 2 1 percent of the drainage basin. 
The close approximation of these proportions, 
despite a considerable discrepancy in estimates of 
the maximum depth (see below), is attributable 
to the steepness of the sides of the enclosing basin. 

The great length and depth that Lake Spring 
attained is consistent with the towering height of 
the closely margining mountains. Except for 
passes, the ridge of the Snake Range to the east 
attains, for most of its length, altitudes higher 
than 2.500 m.. exceeds 3.000 m. in several peaks, 
and reaches 13,063 feet (3,982 m.) at Wheeler 
Peak. Most of the Schell Creek Range, to the 
west, surpasses the altitude of 2.700 m. 

The evidence that this entire valley has no na- 
tive fish ( see below ) confirms the geomorphologi- 
cal indication that the basin is now and probably 
has long been wholly endorheic. 

The hydrography of the basin has been treated 
and the pluvial lake mapped by Rush and Ka/mi 
( 1965). 

Shori^.linfs \nd Inti rior Drain ACii-.. 

We have estimated the maximum depth of Lake 
Spring as approximately 95 m.. on the basis of 
evidence citei.1 below. This estimate is ciinsider- 
ably higher than those presented by C. T. Snyder 
and associates. We have relied on indications of 
higher shorelines that are less bold and less com- 
pletely certain than the ones on which they re- 
lied. Snyder et al. (1964) estimated the maxi- 
mum depth as 265 feet (81 m.). Snyder and 
Langbein (1962. p. 2385) stated that "Spring 
Valley was filled to a depth of nearly 250 feet 
[76 m.| by its Pleistocene lake" and gave figures 
of 5,780 feet for the altitude of "the uppermost 
average stand of this lake, clearly marked by 
shoreline terraces" and of 5535 feet for "the 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



63 



average altitude of the floor." a difference of 245 
feet (75 m.). 

We are of the opinion that the lake rose, prob- 
ably intermittently, to considerably higher levels 
than 75 m., presumably without very long still- 
stands that would have produced strong shoreline 
features — but not to heights approaching the alti- 
tude of the gaps to the southward. The lake out- 
line shown by Snyder cl til. ( 1964) corresponds 
closely with the 5,800-foot (1,768-m.) contour 
on the Ely and Lund 1 : 2 50,000 maps. The Lund 
map depicts that contour forming a very tight 
loop about 3 km. long extending transversely 
across more than half of "Baking Powder Flat" 
in the southern expansion of the old lake floor, 
just short of the southern border of T. 13 N., R. 
67 E. The dikelike structure, followed by High- 
line Road, we verified as a lake bar, which pre- 
sumably was formed under water. Another bar 
is indicated by a tightly closed loop, 4.5 km. 
long, of the same contour, across the lake bed 
near its north end ( shown on the Ely map in T. 
20 N., R. 67 E. ), and below this a complex of 
bars and dunes representing recessional levels of 
the lake. Gravel ridges trending north-south 
along the east shore near Worthington Spring, 
about opposite the major west-east bar of the 
southern basin, were estimated by altimeter read- 
ings, checked against the altitude of 5,878 feet 
near Shoshone entered on the Lund map, to be 
very close to the altitude of 5,845 feet ( 1 ,782 m. ), 
95 m. higher than the 5.535 feet (1,687 m.) 
given by Snyder and Langbein for the altitude of 
the old lake bed (the Ely map shows "5536" on 
White Lake, the present sump). Apparent high 
shoreline features on the east side of the valley, 
extending southward from near the base of the 
bajada just north of where U. S. Highway 6 and 
50 turn southward, were judged when seen on 
July 5, 1959 to be at about the same altitude as 
the gravel ridges near Worthington Spring. The 
uppermost shoreline features are not boldly de- 
veloped, nor would we expect them to be in the 
basin of a lake that did not discharge. Some- 



what lower terraces are undeniable. Below the 
clearly marked feature at 5,780 feet Snyder and 
Langbein ( 1962) indicated six shoreline terraces. 

By far the lowest passes around the rim of 
Spring Valley are three gentle divides near the 
south end of the basin, two into the basin of pluvial 
Lake Carpenter (pp. 65-66) and one into 
Snake Valley of the Lake Bonneville system. All 
three passes are close to the same altitude, which 
is well above any indicated level of Lake Spring. 
One of the two gentle passes between the Spring 
and Carpenter basins, the one followed by U. S. 
Highway 93 leading into the north end of Lake 
( Duck ) Valley, is shown on the Lund 1 : 250,000 
map as being not quite closed by the 6.200-foot 
( 1,890-m.) contours. In the other pass between 
these two basins, leading toward the southern part 
of Lake Valley, the same contours barely cross 
over. The divide into Snake Valley is probably 
at almost the identical altitude, for although the 
same contours here cross the divide farther apart, 
the pass is extremely flat, as we observed in 1938. 
There are no obvious indications of a discharge 
into either Lake Valley or Spring Valley. 

There are no indications that at any time, at 
least not during a late pluvial period. Lake Spring 
ever received any inflow of water from any adja- 
cent basin or that it ever attained a discharge, 
either surface or subterranean. The passes are 
all too high for any surface exchange. As Snyder 
and Langbein (1962) indicated, the floor of 
Spring Valley is lower than that of any surround- 
ing basin other than Snake Valley to the east, and 
the paucity of major springs argues against an 
underground outflow in that direction. The basin 
seems to be completely endorheic. Accepting this 
view, Snyder and Langbein concluded that their 
"studies of the hydrographic regimen that would 
be required to maintain a Pleistocene lake indicate 
that relatively small adjustments of evaporation 
and precipitation would be necessary. The most 
likely combination of adjustments among tho.se 
possible suggests a 30 percent decrease in evapora- 



64 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



tion rates and an 8-inch increase in precipitation." 

(Sec also p. 9. ) 

Remnant Waters. 

White Lal^e. the shallow sump of the northern 
part of the Lake Spring basin, ordinarily holds 
some water, as it did in 193S. when Bert Robison 
and other informed residents ttild us that it had 
gone completely dry in the great drought of 1934. 
for the first time in many years. We were further 
told that in 1934 the valley became so dry that 
cattle had to be shipped out. Several of the trout 
creeks in the Schell Creek Range went dry. Even 
the largest one, Cleve Creek, failed in August to 
reach Cleveland Ranch in the valley. Only a few 
of the deepest of the many spring pools on the 
west side of the northern half of the valley retained 
water. 

A considerable number of trout streams, in- 
correctly shown as wholly intermittent on the 
Ely and Lund I : 250.000 maps, flow to or toward 
the bed of Spring Valley from the Schell Creek 
and Snake ranges. Spring Valley Creek, coursing 
down the narrow northern arm of the valley, 
maintains a slight flow in the dry season and did 
not completely disappear in 1 934. Other streams, 
on the arid alluvial slopes around both northern 
and southern ends of the valley, are intermittent. 

In normal years, many spring pools, ponds, 
sloughs, marshes, and meadows exist on the west 
side of the northern half of the valley, and a spring 
is mapped on the west margin of Baking Powder 
Flat, to the south. The only extensive springs on 
the east valley edge are those on the Shoshone, 
originally Swallow, Ranch, near the base of the 
Snake Range south of Wheeler Peak. Here, in 
1 938. innumerable springs rose in an are about 
5 km. long. These and a few other springs feed 
many ditches and sloughs to form "The Seep" on 
Baking Powder Flat. Other ranches on the east 
side of the valley were said to rely almost en- 
tirely on mountain-stream water. Worthington 
Spring, near the north end of the southern expan- 
sion of the valley, on the east side, had already 
been piped in 1938. 



Fish Life. 

In our earlier report (Hubbs and Miller, 1948b, 
pp. 56-57 ) we mentioned taking in Spring Valley 
two native fishes, "a new sucker, with characters 
that line it up best with a Bonneville, or possibly 
a Colorado species," and the dace we have named 
Rclictiis solilaiins. These we had taken only in a 
spring pool in the course of Spring Valley Creek, 
beside the Stone House ( at the road junction 
west of center of T. 22 N., R. 66 E. ). on the 
old Overland Mail Route, later the Lincoln 
Highway. We have since come to regard 
both species as having been introduced into 
the valley, probably at this old ranch. The 
evidence is presented in the species accounts for 
Catostomiis (Pdiiloslciis) pidtyrhyncluis on p. 229 
and for Rclicius solitaiius on p. 233. Another 
cyprinid, the Utah chub, Gila alraria, which we 
found to be well established in Shoshone Springs, 
we also regard as having been introduced, on the 
basis of testimony that it was brought in by 
Mormon settlers (p. 231 ). The cutthroat trout, 
Salmo clarkii, unde.scribed subspecies, that occurs 
in the mountain streams has also been indicated 
to have been intriKluced (p. 237. and Miller and 
Alcorn, 1946, pp. 177-178). Other species, un- 
questionably exotic, have been introduced. 

PLUVIAL LAKES REGARDED AS HAVING 

BEEN TRIBUTARY TO COLORADO 

RIVER SYSTEM 

We (Hubbs and Miller, 1948b, pp. 55, 96- 
100, 154, 164-165. figs. 25-29) have already 
regarded several basins, each with a pluvial lake, 
as disrupted parts of the pluvial White and Car- 
penter river divisions of the Colorado River sy.s- 
tem. Along with lakes Coal. Bristol, Delamar, 
and Carpenter, we should have included Lake 
Jake ( pp. 57-58. 151. 1 59 ). but not the "lake in 
Long Valley" ( p. 57 ) , namely Lake Hubbs, which 
we now know to have been less disrupted from 
the basin of Lake Newark of the Lahontan 
complex (pp. 26-29). Among these various 
ancient waters, regardable as disconnectctl parts 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



65 



of the Colorado River system, only two. Lake 
Carpenter and Lake Jake, are here treated in any 
detail. 

Physiographic infonnation now available on 
the Lund and Caliente 1:250,000 maps and on 
the 1 965 United States Geological Survey 1 : 
500,000 map of Nevada, prompt us to enter a few 
remarks on some of the other basins. The basins 
of lakes Coal, Bristol, and Delamar, and the de- 
pression of Cave Valley, which Snyder et al. 
( 1964) indicated to have contained a lake they 
named Cave, are now confirmed as being physio- 
graphically most closely related to the drainage of 
the pluvial White River system. However, the 
alluvial deposit in Pahranagat Valley that formed 
Maynard Lake ( Hubbs and Miller. 1948b, p. 98, 
fig. 26) now appears less likely to have resulted 
from the ancient discharge of Lake Delamar, as 
we thought, than from an outflow from a canyon 
immediately south of Delamar Valley. The basin 
that contained Lake Delamar is essentially con- 
tinuous with Dry Lake Valley, the site of pluvial 
Lake Bristol. There is a strong possibility, but 
as yet no definite evidence, that some or all of 
these lakes, all at most shallow, actually main- 
tained a surface discharge into White River 
waters. The same holds true for the irregular 
chain of lake-containing valleys (p. 35) that 
extend between Railroad Valley and pluvial White 
River and may mark the course of a very remote 
discharge of Lake Railroad. 

Pluvial Lake Carpenter 

Drainage basin in extreme southern White Pine 
County and, chiefly, in Lincoln County, eastern 
Nevada (fig. 1 ). 

We proposed (Hubbs and Miller, 1948b. pp. 
98-100) for this ancient body of water the name 
Lake Carpenter, which has been accepted by 
Snyder et al. (1964) and by Feth (1964). Like 
most of the other depressed basins, it is repre- 
sented at present by an extensive alluvial flat, 
that of Lake ( or Duck ) Valley. 

The drainage basin of this ancient lake was 



bounded on the short northwestern sector by the 
Lake Steptoe basin and on a short northeastern 
sector by the Lake Spring basin; on the east side 
chiefly by the drainage of Lake Spring and very 
slightly by the Lake Bonneville watershed; 
around the south end by tributaries of what we 
have called pluvial Carpenter River (the pre- 
sumed outlet of Lake Carpenter), a tributary to 
the pluvial White River and Colorado River sys- 
tems: and on the west side by headwaters of 
pluvial White River and by affluents of Lake 
Cave, which probably discharged into White 
River. 

The drainage basin and the lake were roughly 
similar in north-south orientation and in form. 
The basin was 72 km. long and. toward its south 
end. 34 km. in maximum width. Corresponding 
greatest dimensions of the pluvial lake were 38 
and 1 1 km. The northern end of the basin and 
of the lake were pointed; the southern end. 
rounded. 

We estimate that at its highest stage Lake 
Carpenter occupied 247 sq. km., or 20 percent of 
the area ( 1.257 st]. km.) of its drainage basin. 
The lake was obviously shallow, for the lowest 
altitude for the lake flat given on the Lund I : 
250,000 map is 5.915 feet (1.803 m.) and the 
outlet was somewhat below the 6,000-foot 
(1.829-m.) contour, indicating the lake depth 
as somewhat less than 26 m. This extensive ac- 
cumulation of water is attributable to the circum- 
stance that the elongate drainage basin was 
closely bounded by mountain ranges rising in 
general more than 600 m. above the lake level. 
and to the altitude of 3.254 m. at one peak. Mt. 
Grafton. 

There was obviously no pluvial inflow of sur- 
face water into the basin of Lake Carpenter, al- 
though there were two low and gentle passes be- 
tween this basin and that of Lake Spring (p. 
63 ) . 

There is some disagreement as to whether Lake 
Carpenter rose high enough to discharge south- 
ward into the course of pluvial Carpenter River, 
an affluent of the Colorado throush the lower 



66 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



pluvial White River. Carpenter (iMI5. p. 44) 
indieated that the lake mi diseharged. He stated: 

During the tiumid Pleistocene epoch this 
lake drained southvsard into Meadow Valley 
through a channel ahout 5(1 I eel deep and 
one-fourth mile wide, this channel begins at 
aboLit the latitude ol Poney Spring and be- 
comes progressively deeper toward the south. 
East of Pioche Range it is about 100 feet deep 
in places and is cut through bedrock. 

Snyder ci al. {\9b4). iiowever, claimed that the 
lake did not spill. Morrison ( 1965, fig. I ) indieated 
the outlet. If the lake did not overflow in late 
pluvial time, it may have done so on some early 
pluvial oceasion(s). Although the passes to the 
Spring and Bonneville watersheds are also low, the 
one to the southward (to Carpenter River) seems 
to be the lowest. It is indieated on the Lund map 
as the lowest, just below the 6.()()0-foot ( I ,S29-m.) 
contours, whereas the passes leading to the Lake 
Spring and Lake Bonneville watersheds are indi- 
cated, respectively, to be a little below and a 
little above the 6,20()-foot (l,89()-m.) contour. 
Present underground discharge through Patter- 
son Wash was indicated by Rush and Eakin 
( 1963). 

Although we took no elevations, we are led 
from our reconnaissance of July, 193S. definitely 
to favor Carpenter's view that l^ake C arpenter 
did discharge to the southward. Significantly, 
local testimony indicated that Wilson Creek, the 
main affluent to the lake bed (see below), in 
times of flood turns southward to join Patterson 
Wash, which definitely is in the southern drain- 
age. Our field notes indicate, further, that there 
may be a low spot south of Gey.ser, but that even 
from there the general trend is southward ( this 
point is not well verified by the few altitudes en- 
tered on the Lund map ) . From near Pony Springs 
(a little south of the pluvial lake as mapped by 
Snyder ct al. and by us ) there are beds and ter- 
races of considerable magnitude, indicating a 
former river system (which would have been the 
head of pluvial Carpenter River). 



RiMNANi Waii;r.s and Fish Life. 

The only extensive spring waters in the Lake 
Carpenter basin are those just south of Geyser 
Ranch, near the north end of the old lake bed. 
These waters discharge through very deep sloughs 
into a scries of small lakes, in a broad area of 
wet marshes. On July S, 19.iS, the water in a 
slough registered 20.0 C. (air, 21.1 ) and con- 
tained considerable submerged and emergent 
vegetation. The only other springs near the lake 
bed are Pony Springs, on the west side near the 
south end. The only stream of any consequence 
ill the basin is Wilson Creek, which flows north- 
westward toward the lake bed from the north 
side of Mount Wilson, the altitude of which is 
mapped as 9.2'-)6 feet (2.833 m.). On the next 
day. this stream, where examined 14..'^ km. east 
of Pony Springs, was about 1 m. wide and 10 cm. 
deep, with a good current of clear water. The 
hydrography of the basin was treated by Rush 
and Eakin ( 1963). 

On the same trip, we found no fish that surely 
appear native in any of the waters just men- 
tioned. Trout" had been planted in Wilson Creek, 
which was posted as closed to fishing. No fish 
were seen in Pony Springs. In a spring-fed ditch 
in the complex of waters near Geyser we collected 
carp. Cyprliiii\ airpio. and L'tah chubs. Gilii 
iilniria ( UMMZ 124790). Local residents were 
too new to know anything abiuit the introduction 
of chubs, but the eliainage relations are such as 
to render it virtually certain that they had been 
introduced. Some testimony indicates stocking 
from Utah (p. 231). Plausible sources nearby 
were the springs at Shoshone, where they were 
almost certainly introduced (p. 64), and Big 
Springs or the adjacent slough south of Garrison, 
Utah, nearby in the Bonneville Basin (a native 
habitat for the species). 

Pluvial Lakl Jaki- 

Drainage basin in southwestern White Pine 
County, east-central Nevada (figs. 7, 13). 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



67 



This, one of the smaller of the enclosed Pleisto- 
cene lakes, was named by us ( 1 948h. pp. 57-58 ) 
Lake Jake, which designation has been accepted 
by Snyder et al. (1964) and Feth (1964). 

The drainage basin is bounded by the water- 
sheds of the following pluvial waters: Lake 
Hubbs to the northwest; Lake Gale very narrowly 
to the northeast; Lake Steptoe to the east; White 
River to the southeast and south; and lakes Rail- 
road and Newark to the west. 

The whole drainage basin is roughly diamond- 
shaped. 51 km. long by 32 km. wide, with a sub- 
central elliptical fault-block depression, Jakes 
Valley, roughly 34 • 17 km., lying between 
mountain fronts. Within the drainage basin, the 
depressed block and the lake are displaced to the 
eastward. 

As mapped by Snyder ci ul. ( 1964). slighth 
larger than we did in our first report ( Hubbs 
and Miller, 1948b. pp. 57-58). Lake Jake mea- 
sured about 7 X 17 miles (1 1.3 X 27.4 km.) 
and covered a stated area of 67 square miles 
( 174 sq. km.). 16 percent of the drainage basin 
of 429 square miles (1,112 sq. km.), and they 
indicated the depth of the non-discharging lake 
as 65 feet (20 m.). Although we have seen the 
basin only from near the north end of the ancient 
lake, we believe that these were underestimates of 
the size of the lake. We draw this conclusion 
from the cursory field observations, local testi- 
mony, and, particularly, from an examination of 
four 15-minutc quadrangles (clockwise from 
northwest; Illipah, Riepetown, Preston Reservoir, 
and Treasure Hill). 

We now compute higher values; the lake as 
measuring 12 X 31 km., covering an area of 249 
sq. km. ( 22 percent of the drainage-basin area of 
1.114 sq. km.), and the lake depth as 150 feet 
(46 m.). Thus, we compute the lake as slightly 
longer. 1.4 times greater in area, and 2.3 times 
deeper. We have arrived at our estimates as fol- 
lows; The altitude of the central depression 
(without, however, a playa, according to local 
testimony) is slightly below 6,295 feet. This is 
the figure given in the Riepetown Quadrangle for 



Jake VABM (Verified Altitude Bench Mark). 
which is near the end of a dry wash. We approxi- 
mate the lowest point as 6.292 feet ( 1,918 m.). 
On reasoning given below, we assume that the 
top level of probably occasional discharge is 
about 6.442 feet ( 1,964 m.), thus providing the 
estimate of 150 feet (46 m.) for the maximum 
depth of the lake. 

A minimum approximation of the depth at 
least approaches 100 feet (30 m.), because the 
Riepetown Quadrangle shows, near the northwest 
corner of the lake bed, the interpolated contour of 
6,380 feet ( 1,945 m. ) in the form of complex 
bars that were presumably formed under water. 
These circumstances indicate a water depth in 
excess of 88 feet (6,380-6,292). or 27 m. In 
line with these data on a minimal size estimate we 
have mapped a second lake outline, as a dashed 
line at the 6,400-foot contour (1.951 m.). yield- 
ing estimates of 108 feet (33 m.) for the lake 
depth, of 26 ■' 1 1 km. for the major dimensions, 
and 186 sq, km. for the lake area (17 percent 
of the drainage-basin area). Other indications 
on the same and other quadrangles bespeak the 
reasonableness of these estimates. Nearby, in the 
axis of the bed of the lake, a labelled "Gravel 
Pit" suggests a beachline. On the east side, an- 
other "Gravel Pit" is shown, beside a verified- 
altitude benchmark at 6.363 feet (1.939 m.). 
Near the south end of the lake bed there is an 
ancient gravel bar (see below), which on the 
Preston Reservoir Quadrangle is designated by 
the suggestive term of "Railroad Crossing," just 
above the 6,360-foot (1,939-m.) contour, and 
this bar was presumably formed under water. 
Furthermore, apparently longshore features are 
mapped on the same contour on the Illipah 
Quadrangle (in Sec. 11. T. 16 N.. R. 59 E. ) 
and on the Preston Reservoir Quadrangle ( near 
the southeastern margin of the old lake). 

Several lines of circumstantial evidence lead 
us to regard as highly plausible the hypothesis 
that Lake Jake occasionally rose to. or at least 
well toward, the assumed outlet level, which lies 
near the north end of Jakes Wash. That lone 



68 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



t'lood-watcr course, in the headwaters of pluvial 
White River, is separated by a very low and 
gentle saddle from a wash, in the same linear de- 
pression, that runs northward onto the bed of 
Lake Jake. One line of evidence is that the basin 
is hemmed in by rather high nnnmtains. the Egan 
Range to the east and the White Pine Range to 
the west. Tributaries forming a rather extensive 
system in the White Pine Mountains still carry 
water, feeding the sizable Illipah Creek (see be- 
low), and these mountains retain snow in winter. 

The percentage of the area of the drainage 
basin that was covered by the lake (as now 
treated), namely 17 to 22 percent, is average for 
the whole area. This area is one where we would 
expect the fluctuations in precipitation to have 
been great. During unusually moist periods the 
lake might well have risen rapidly to or at least 
well toward the sill. The lack of a deep trench 
in Jakes Wash, however, is evidence that an out- 
let if attained was not greatly prolonged, or was 
so ancient that the channel has tieen obliterated 
by lateral aggradation. 

Information regarding this basin, in part sum- 
marized by us ( Hubbs and Miller, iy48b. p. 56). 
was obtained on June 26. 1942. from Mr. Moor- 
man, the only rancher in the drainage basin (the 
Moorman Ranch is also mapped as Illipah, and 
the mountains margining the lake basin on the 
west form the Moorman Ridge). Jakes Valley, 
we have heard, was named for "Dutch Jake," 
the earliest settler at Illipah. Moorman's father 
came to Nevada in the l<S60's and settled at 
Illipah in 1S96. Mr. Moorman informed us that 
the valley has no dry lake bed. but to the south 
there is an oval white-sage area about 1 .5 miles 
( 24 km. ) long, surrounded by lines of water 
levels, indicating successive recessions of the 
former lake. Across this, he said, is a broad bar 
of round stones somewhat cemented together, 
which we assume to be an ancient gravel bar, the 
"Railroad Crossing" (mentioned alxive). The bar 
had been broken in one place by a washout, which 
had been filled in to make a reservoir ( shown on 
the Preston Reservoir Ouadranszle ) for stock. 



Remnant Wathrs and Fish Life. 

Mr. Moorman and ranch hands informed us 
that Illipah Creek is the only permanent stream 
in the basin, and reported its flow in the summer 
as 3-4 second-feet, all of which was used at the 
ranch. On June 26. 1942, at the mouth of the 
canyon about I km. above the ranch, the rather 
muddy, perhaps somewhat swollen stream, was 
about 3 m. wide and flowed over a stony bottom. 
Mr. Moorman said that the main creek ( Halstead 
and Cottonwood creeks are tributaries) rises in a 
large permanent mountain spring. His further 
testimony was recounted by Miller and Alcorn 
(1946, pp. 177-17.S). Several other mountain 
springs are mapped at the headwaters of Illipah 
Creek, and on the south side of the northern 
divide. 

According to Leigh ( 1964, p. 61 ). "Illipah 
Spring in Momoke Hill, near Little Antelope 
Summit, is a splendid spring gushing from a cleft 
in solid rock 2,()()(),()()0 gallons of water each 
day." 

Mr. Moorman was positive that there are no 
native fish in Illipah Creek, or anywhere else in 
the basin. His testimony regarding the stocking 
of trout and other data on the introduction of 
trout into the creek was recounted by Miller and 
Alcorn. Mr. Moorman's statements we regarded 
as highly reliable. He well knew the minnows 
and suckers of White River to the south, where 
a rancher "used to fry the minnows in the round 
to eat." He nincd also the use of such native fish 
for "coyote scent" and knew of miniTOws in vari- 
ous other waters where we had found them. 

PLUVIAL LAKE PINE ( "WAH WAH"). 
WITHIN BONNEVILLE SYSTEM 

The vast hydrographic basin of Pleistocene 
Lake Bonneville, which, among the many divi- 
sions of the Great Basin, was approached in 
magnitude only by that of Lake Lahontan ( in 
terms both of basin and lake), was essentially 
integrated, in strong contrast to the carved-up 
area we have Luidei' primary treatment. As the 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



69 



great inland Bonneville sea desiccated, some 
basins, as those of Utah Lake and Sevier Lake, 
became independent. In addition, a few tributary 
lake basins of fault-block origin, charted by Jones 
(1940) and mentioned by us (Hubbs and Miller, 
1948b, pp. 32. 148, 156), existed peripherally 
to Lake Bonneville at the top (Bonneville) level. 
One of them, though wholly surrounded by inte- 
grated parts of the Bonneville system, was rela- 
tively near the enclosed basins that we are now 
discussing, and has been included in the study. 

It is possible that at some time more remote 
than we are emphasizing, some other basins may 
have been included in the Bonneville watershed, 
for the intervening passes are lower than those 
into other depressed basins. These are the 
Waring-Steptoe system and the basin of Lake 
Spring (see above). 

The drainage basin of Lake Pine, that of Pine 
( = White Sage ) Valley, lies in southwestern 
Millard County, in western Beaver County, and, 
very slightly, in northwestern Iron County, south- 
western Utah ( fig. I ) . 

Lake Pine was treated and named as a pluvial 
as well as a modern body of water by Jones 
( 1940), as we noted in our brief treatment of 
the lake (Hubbs and Miller, 1948b, p. 32). The 
lake was renamed "Wah Wah" by Snyder et al. 
( 1964). We prefer to retain the name Lake 
Pine on the basis of priority and because Wah 
Wah Valley, Wah Wah Springs, Wah Wah Field 
Station, and Wah Wah Dry Lake all lie to the 
eastward, beyond the Wah Wah Mountains. 

Pluvial Lake Pine was unique among those 
herein treated in having been completely sur- 
rounded by the drainage basin of Lake Bonne- 
ville. 

The drainage basin, as delineated from the 
Richfield 1 :250.000 map and the Wah Wah Sum- 
mit 15-minute Quadrangle, is 75 km. in greatest 
length (north-south) and 39 km. in maximum 
width, north of the middle. The basin is largely 
surrounded by the Needle and Wah Wah moun- 
tains, which converge southward, and an inter- 
polated mountain at the north, between two rather 



gentle passes. The dimensions of the lake, as 
traced by us. were 15 km. long and, near the 
north end, 10 km. in maximum width. 

Areal dimensions of lake and basin were given 
by Snyder cl al. ( 1964) as 41 and 720 square 
miles, indicating that the lake covered 6 percent 
of the area of the drainage basin. Our essentially 
similar numbers are 102 and 1,912 sq. km. (5 
percent ) . 

This relatively limited accumulation of water 
may be attributed to the moderate height of the 
surrounding mountains, the rather low and broad 
passes to the north, the position in the rain shadow 
of the lofty Snake Range, and the southern loca- 
tion. Sharp southward decrease in surface-water 
accumulation in pluvial time, matching the pres- 
ent southward increase in aridity, has been indi- 
cated in the several general treatments of pluvial 
waters in the West. However, the basin still con- 
tains a live stream (see below). 

The topographic maps now available, as well 
as the field reconnaissance, show clearly that the 
low passes between this wholly endorheic basin 
and the Snake Valley arm of Lake Bonneville 
were far above any possible lake levels, and the 
pass toward Escalante Desert was even higher. 
The doubts we expressed in 1948 (p. 32). when 
topographic data were not available, were un- 
founded. There are no suggestions of any inflow 
of surface water from any other basin. 

These physiographic relations became evident 
to us in a reconnaissance of the basin on June 24. 
1 950. We found that there are definite indications 
of a beachline, and now note that these are ap- 
proxin:ately on the enclosed 5.200-foot (1.585- 
m.) contour on the topographic maps — the line 
that Snyder et al. ( 1 964 ) apparently followed in 
charting the lake. Along the road from state 
highway 2 1 northerly toward the headquarters 
of the Desert Range Experimental Farm, a shore- 
line was suggested at 2.5 km. where the road is 
now mapped beside the 5.200-foot contour. At 
4.0 km., at the gates to the Experimental Farm, 
still beside that contour, we encountered a sharp 
slope that seemed to surround the basin, at least 



70 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



on the southwest side. At the Farm we obtained 
confirmatory evidence from Selar S. Mulchings 
and other staff members. They had ai.so con- 
cluded that the top beachlines are approximately 
at the entrance gates. They had found tufa on 
rock along the lake terraces and showed us sam- 
ples. These were taken at aboLit the same level 
as the entrance gates, at places estimated to be 
about 2 miles from the playa and 1 00 feet higher 
( the depth of the lake was presumably nearer 200 
feet = 60 m.. for the top beachlines approximate 
5,200 feet and an elevation of 3,0(S6 feet is shown 
outside the playa). They mentioned further that 
clean gravel had been dug by the corral of the 
abandoned ranch near the 5,20()-foot contour on 
the south side (such gravel is commonly of shore- 
line oriiiin). 



Remnant Waters and Fish Life. 

The topographic maps show an ephemeral 
pond at the east end of the playa. The longest 
tributary, from the south, was found to be dry. 
but in T. 27 S.. on the line of R. 16-17 W.. the 
terrace-flanked valley, about 1 km. broad, indi- 
cated greater pluvial flow. At the time of our 
visit (June 24, 1^30), an ample stream was flow- 
ing in the well wooded tributary Pine Grove 
Canyon, in T. 27 S., R. 16 W. (the place where 
pine lumber was said to have been obtained for 
the famous organ pipes of the Tabernacle in Salt 
Lake City). Where examined near its lower end 
in the nunilli of the canyon, this stream was fish- 
less, and we were told in 1^50 by Otto Fife, a 
well informed naturalist at Beryl Junction, Utah, 
that the trout m Pine Grove Creek had been 
introduced. 



FISH LIFE OF BASINS IN NORTH-CENTRAL GREAT BASIN 



As indicated in i)ur earlier sLunmary ( Hubbs 
and Miller. l94Nb), correlating the hydrographic 
history and the residual fish faunas of arid west- 
ern NtMth America, four native fish species occur 
in the north-central part of the Great Basin herein 
under more detailed treatment. Their respective 
occurrences in the isolated and limited waters of 
the several basins are mentioned in the preceding 
discussions of these basins, and are outlined in 
tables 2 and 3. 

GENERAL TREATMENT 

Fish Fauna of Area: Its Hic.h Endemism and 
E.XTREME Depauperation 

It is indeed noteworthy that only four native 
fish species have survived ui the largely isolated 
springs in the north-central part of the Great Basin 
under special treatment, although the area in- 
volved embraces 54,028 sl\. km. and is surrounded 
by the Lahontan. Columbia, and Coloradii drain- 



age basins, each of which is endowed with a much 
less depauperate fish fauna ( Miller, 1958 ). These 
four indigenous species, however, include two 
relicts wholly endemic to the area, as well as 
representatives of twd wide-ranging Western 
species, each much fragmented by local differ- 
entiation, which we interpret as being in part on the 
subspecies level. Such endemism is often char- 
acteristic of fishes that inhabit springs, even m 
areas where the faima is much more profuse (as 
Armstrong and Williams ( I 97 I ) have noted for 
the Tennessee River system). 

One of the luc endennc species, the relict dace 
{ Ri'liciiis M'lluirius. pp. 196-226), representing 
also a genus of Cyprinidae confined to the area, 
occurs native only in two contiguous pairs of 
basins, each compiising a separate endorheic 
pluvial drainage system — that of lakes Franklin 
and Gale and that of lakes Waring ( Lake Ante- 
lope excluded ) and Steptoe ( Upper Lake Steptoe 
excluded). This species occurs alone in these 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



71 



Table 3. Occurrences of ncilive fislies in 21 phiv'ud-lake basins in the north-central Great Basin. 





Rhinichthys 




Gila 




Reliciii.\ 


Crenichthys 


Pluvial lake basins' 


OSClllllS 




bicolor 




solilariiis 


nevadae 


Basins related to Lake Lahontan 














L. Gilbert R. 


o. rcliqutis 




— 




— 


— 


L. Diamond R. 


o. rohiislmr 


G. 


b. obesa'' 




— 


— 


L. Diana 


— 




— 




— 


— 


L. Yahoo 


— 




— 




— 


— 


L. Newark 


— 


G. 
G. 


/>. newarl<eiisis 
b. ciichiln 




— 


— 


L. Hubbs 


— 




— 




— 


— 


L. Clover R. 


(). oligoporiis 


G. 


b. isoiiilii 




— . 


— 


R. 


(>. leltioporiis 












Basins related to L. Lahontan and 














Colorado R. 














L. Railroad 


— 


G. 


b. subspecies' 




— 


C. nevailac 


Little Fish Lakes 


— 


G. 


/'. subspecies 




— 


— 


L. Lunar Crater 


— 




— 




— 


— 


L. Snyder 


— 




— 




— 


— 


Basins between Lahontan and Bonne- 














ville basins 














L. Franklin 


— 




— 


R. 


soUturiits 


— 


L. Gale 


— 




— 


R. 


soHtariiis 


— 


L. Waring 


— 




— 


R. 


solitarius 


— 


L. Steptoe 


— 




— 


R. 


solilariiis 


— 


Upper Lake Steptoe 


— 




— 




— 


— 


L. Antelope 


— 




— 




— 


— 


Basin related to Bonneville and Colo- 














rado systems 














L. Spring 


— 




— 




— 


— 


Basins tributary to Colorado River 














L. Carpenter 


— 




— 




— 


— 


L. Jake 


— 




— 




— 


— 


Basin surrounded by Bonneville 














system 














L. Pine 


— 




— 




— 


— 



^ Tributary relations siiown by indentations. 

- Slightly modified endemic races referred to R. oscitlus rubiistus. 
3 Strongly modified endemic races referred to G. bicolor obesa. 
* Several subspecies, ail iindescribed. 



two systems, which together conipri.se an area of 
14,682 sq. km. (27 percent of the total study 
area). The many local populations of the relict 
dace in these systems, along with a few stocks in 
the basin of Lake Spring into which it was intro- 
duced, have been subjected to intensive analysis, 
to document the relict nature of the genus and 
species and its limited local variation. Together 
with the drainage basin of Lake Clover, the Frank- 
lin and Waring systems form the northeastern 
sector of the study area. 

The other relict species. Crenichthys nevadae 
Hubbs (1932), represents a remnant genus (p. 
227 ) of Cyprinodontidae with two fully distinct 



species, C. nevadae (p. 227). confined to Rail- 
road Valley, the sump basin of the pluvial Lake 
Railroad (pp. 32-36) drainage basin, and C. 
baileyi (Gilbert), a relict endemic in the now 
isolated remnant waters of pluvial White River, 
which during the pluvial period(s) was tributary 
to the Colorado River (Hubbs and Miller, 1941, 
pp. 2-3 ) . Both species are confined to hot springs 
in Nevada in the northernmost extension of the 
range of the Cyprinodontidae in western North 
America (Miller. 1958, pp. 203-207, fig. 15). 
The detailed systematic treatment of Creuiclithys 
is deferred, but annotated synonymies of the genus 
and of C. nevadae are included (pp. 227-228), 



72 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



along with a list of tlic known specimens of that 
species. 

All other native fish of the study area are min- 
nows that we refer to two species. Rliinichrhys 
osciihi.s and Gilii iSiphati'li's) hicolor. each of 
which is widespread through western United 
Slates and is subject to almost endless local dif- 
ferentiation. All of the local populations within 
the area, of both species, have been subjected to 
extensive analysis, particularly to document the 
marketl local differentiatitin, even within a single 
endoiheic basin, that often has accompanied their 
isolation in the remnant spring waters. For both 
species the variational analysis has been extended 
to include populations from the Humboldt River 
system, taken to represent an approach to the 
presumed ancestral stock from which the deviating 
populations of the .several basins arose. The 
basins that have retained remnants of Rliinichthys 
and Gila are those, along the northern edge of the 
pluvial-lake area of the Great Basin, that dis- 
charged in late pluvial time, or earlier, into Hum- 
boldt River of the Lake Lahontan drainage sys- 
tem. These basins, from west to east (fig. 1 ). are 
those of lakes Gilbert. Diamond. Newark, and 
Clover. The basins of lakes Diamond and Clover 
have retained both genera, that of Lake Gilbert 
retained only Rhinichthys ( until its very recent 
apparent extinction), and that of Lake Newark 
has maintained only GiUi. 

That the Rhinichthys osciihi.s and the Gila 
hicolor populations in the northern basins under 
treatment stemmed from the populations in the 
Humboldt headwaters seems obvious for a num- 
hcv of reasons. The physiographic evidence is 
strong and definite. Both species are ubiquitous 
in those headwaters, and almost certainly have 
been there through at least late Pleistocene time. 
Both species seem preadapted to survive in iso- 
lated waters. The characters of the isolated forms 
are plausibly derivable from those exhibited by 
the Humboldt headwater populations of Rhiiiicli- 
thys osciiliis rohiistiis and Gila hicolor ohcsti ( pp. 
106-113, 151-153). Particularly significant is 
the circumstance that in various characters the 



Humboldt headwater races of Gila hicolor ohc.sa 
very definitely approach the presumably derived 
forms occurring in the basins of pluvial lakes 
Diamond, Newark, and Clover, as is stressed in 
the systematic accounts (pp. 153-1X0). 

The only other pluvial lake basin among the 
21 herein treated that has retained twci fish 
species is that of Lake Railroad, which contains, 
in addition to Crcnichthys ncviidac. as we have 
noted ( Hubbs and Miller, 1948b, p. 91 ), a chain 
of differentiated popiilations of Gila hicolor, the 
detailed analysis of which is scheduled for sub- 
sequent publication. The Gila hicolor popula- 
tions in the Lake Railroad basin (including the 
Little Fish Lakes subdivision) seemingly were 
derived from the basin of pluvial Lake Newark, 
by passage over the low divide (p. 35). as we 
have surmised ( Hubbs and Miller, 1948b, p. 91 ). 
The only other Gila hicolor populations that have 
survived south and cast of the Humboldt River 
system are those of Dixie Valley, the basin of 
pluvial Lake Dixie, and those of Big Smoky 
Valley, the basin of pluvial Lake Toiyabe. We 
believe (p. 14) that the Toiyabe population of 
Gila hicolor was similarly derived, along with 
Rhinichthys osculiis, from the basin of pluvial 
Lake Gilbert. Cursory examination of the form 
of Rhinichthys in Big Smoky Valley leads us now 
to think that it warrants subspecific separation, 
though it definitely appears to have been derived 
from the Lahontan foi in. R. o. rohustus. We have 
already indicated (Hubbs and Miller. 1948b, p. 
44 ) that the Gila hicolor populations of the Lake 
Toiyabe basin appear to represent two endemic 
subspecies. These are yet to be described. The 
local representative of R. o. rohiistiis, a subspecies 
in our opinion, has recently been described as a 
new species by Lugaski ( 1972), 

To summari/.e. the finu' fishes that are indige- 
nous to the 2 1 pluvial lake basins under discussion 
comprise two basic categories, which are strictly 
allopatric. except for limited sympatry in the 
large basin of pluvial Lake Railroad. These two 
categories, each made up of two species, are as 
follows: 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



73 



(A) Fishes of widespread type that, in the north- 
central Great Basin, appear to have stemmed 
from past stream discharge into the major 
drainage basin of phivial Lake Lahontan, through 
the Humboldt River system. These comprise two 
chains of local forms. The members of each group 
are treated as subspecies of a separate cyprinid 
species: Rhinichthys osculiis, in the basins of 
pluvial lakes Gilbert, Diamond, and Clover, and 
Gild hicolor, in the basins of pluvial lakes Dia- 
mond, Newark. Clover, and Railroad (including 
the Little Fish Lakes subdivision). 

(B) Two fishes of localized distribution, a 
cyprinid, Relictiis soUtarius, whose origin is not 
apparent, and a cyprinodont, Creniclithys iieva- 
dac. whose origin seems to date from some rela- 
tively very ancient discharge into the Colorado 
River system. The cyprinid is confined as native 
to the basin of pluvial lakes Franklin and Gale, 
and to the basin of lakes Waring and Steptoe. The 
cyprinodont is restricted to the central basin of the 
pluvial Lake Railroad drainage system. 

The last two species listed, as noted before, 
appear to be old relicts, whose congeners are 
presumably to be found in the fossil record. Some 
information has appeared on the fossil history of 
Empetrichlliys, the one near relative of Creni- 
clithys ( Uyeno and Miller, 1 962 ) , but nothing has 
been discovered regarding the paleontology of 
Relictiis. In fact, no trace of Cenozoic fossil 
fishes has been unearthed in any of the 21 pluvial 
lake basins under treatment: a great pity. Such 
evidence should be diligently sought. A very im- 
pressive demonstration of light being thrown on 
the knowledge of the fish life in the Great Basin 
through paleoichthyological studies was accom- 
plished by recent research on fossil fishes in the 
Lake Bonneville deposits ( announced by Stokes, 
Smith, and Horn, 1964 and described by Smith, 
Stokes, and Horn, 1968). Recent expanding at- 
tention to the Cenozoic paleoichthyology of the 
West is very propitious. 

Of the 21 pluvial-lake basins herein treated, 1 1 
are now totally devoid of native fish life. 7 have 



retained one species (recently extinct in one 
basin), and only 3, as is mentioned above, have 
retained 2 species ( table 3 ) . 

The faunal deficiency of the north-central 
Great Basin, throughout the drainage basins of 
the 21 pluvial lakes under current treatment, is 
exemplified not only by the great paucity of 
species retained in any of the endorheic depres- 
sions and the complete absence of native fishes 
in many of the 21 basins, but also by the lack of 
various taxa that would likely have persisted in 
the area if it had retained the profuse array of 
lakes and streams that existed in pluvial time. 
Such circumstances more than offset the retention 
within the area of two relicts: of Relictiis solitariiis 
in the Gale-Franklin and Steptoe-Waring drain- 
age basins, and of Creniclithys nevadue in the 
Lake Railroad basin. 

It is almost certain that native salmonoids are 
lacking, although subspecies of Salmo chirkii 
occur in surrounding basins. Their absence in 
the area being reported upon seems attributable 
to the very small size of the cooler mountain 
streams, from which native fish would presum- 
ably have been eliminated by the rare torrential 
floods. In addition, the streams of the Ruby and 
East Humboldt ranges, in the Franklin and 
Clover basins, were, it may be assumed, largely 
eliminated by the montane glaciation there (Sharp. 
1938). Mountain whitefish of the genus Pro- 
sopiitm presumably occurred in the study area, 
for they have survived in the Bonneville and 
Lahontan systems to near their southern limits. 

The families, genera, and species that failed 
to survive generally inhabit permanent waters of 
some magnitude, most commonly streams that 
persist onto the valleys, where they can resist 
lethal flushing by flash floods. In the basins 
under study they presumably succumbed to in- 
creasing aridity, because they were less pre- 
adapted than Rhinichthys osciiltis and Gilo 
hicolor to the gross deterioration of surface waters 
that characterizes the north-central Great Basin. 

The depauperation of the fish fauna of the 
north-central Great Basin is further emphasized 



74 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



and paiticulaiizcd when attention is drawn to the 
consideiable number of now extirpated families 
(Salmonidae, C'oiegonidae, Catostomidae, and 
C'ottidae). of various genera and species, that 
ahiiost surely abounded during some pluvial 
periods in at least some of the 21 basins under 
study. On the basis of their profuse retention in 
the adjoining Lahontan and Bonneville com- 
plexes (Synder. I 9 1 7; Hubbs and Miller. 1948b; 
Miller. 1962; La Rivers. 1962), and their sur- 
vival in some of the adjoining basins, it is plausible 
to postulate that during pluvial times various other 
species occurred in two sets of drainage basins 
among the 21 treated. 

It may be assumed that the basins that dis- 
ciiarged into Lake Lahontan via Humboldt River 
(the basins of lakes Gilbert, Diamond, Newark, 
and Clover) harbored, in addition to the trout 
Sdlnio chirkii and the whitcfish Piosopiiiiu 
willidiiisoni (Girard); suckers. CdlostoDiits iCatos- 
toinii.s) ttiliocnsis Gill and Jordan and C. {Paiilo- 
slciis ) pliiiyihynchiis, and probably Cluismistes 
ciijiis Cope; another minnow. Richanlsoiiius 
c>i>rciiius (Girard). and a sculpin. Coitus hcldingi 
Eigenmann and Eigenmann. Similarly, the basins 
that are thought to have discharged into Colorado 
River ( the basins of lakes Carpenter and Jake and 
probably Railroad ) presumably harbored Colo- 
rado River endemics, including suckers of the 
subgenera Cufostonius and Piinloslcus of the 
genus Ccitosloniiis. and the genus Xyidinlicir. 
probably minnows of the tribe Plagopterini ( Mil- 
ler and Hubbs, I960) and of the subgenus Gila; 
perhaps still other cyprinids; and perhaps also 
salmonoids { Prosophmi and Salnio). Extirpation 
must have been the rule. 

The concept of the extreme depauperation of 
the fish fauna in the many smaller, largely en- 
dorlieic basins of the Great Basin is accentuated 
when attention is directed to the southern part of 
the Great Basin, where conditions both in pluvial 
and postpluvial time have been more arid than 
farther north. The entire area of the Great Basin 
south and southwest of the region under treat- 
ment, and south and southeast of the Lahontan 



system, excluding only the less depauperated 
Death Valley system (Miller, 1946. 1948) en- 
compasses, as delineated and measured by Snyder 
el al. (1964): 

One pluvial lake basin, loisabe. of 3..Vs4 sq. km., 
with two lish genera and species ( Rhinichrlns 
nscullis and iiilti /)/( n/i'/', p. 72); 

Two pluvial lake basins, l^ixie and C'oliinibus, of 
6,314 anti 3,<i2(i sq. kni.. respectively, eacti with 
a single speeies. (iiUi hu<>lor (and the Dixie basin 
includes the lishless l.abon section, of 777 sq. km., 
and this species in the Columbus basin is confined 
to the Fish lake tributary section, of 2.779 sq. 
km.); 

A total of 5 1 pluvial-lake basins, as recognized by 
Sn\'der <■/ iiL, comprising 42 drainage systems, 
occupying a total of 67.7.^7 sq. km., all wholly 
devoid of an\ native fish life. 

Of these 54 basins, the 3 that have retained fish 
obviously derived these from the Lahontan sy.s- 
tem, and some of the .^ 1 now fishless basins pre- 
sumably also had some connection with the 
Lahontan system. 

Except for the Death Valley system ( the pluvial 
Lake Manly complex), the drainage systems that 
have retained no fish into modern times con- 
stitutes 84 percent of the total vast area and 82 
percent of the total number of drainage systems 
included. 

The fish fauna of the Lake Manly complex 
has been treated h\ us ( Miller. 1948; Hubbs and 
Miller, l94Sb, pp. 77-SS). 

Rl-A-SONS 1 OR im: DliPAUPliRAHON OF THE 

Fauna, with SoMii CouNTERACTtNc; Differ- 
entiation. 

Why only four fish species have survived in 
the basins of the 21 pluvial lakes under special 
consideration, and in only a few of the residual 
bits of water, and why these particular species 
have persisted, call for consideration. Plausible 
reasons are in part zoogeographical, dependent 
on paleohydrographic history, and in part ecologi- 
cal and physiological, dependent on adaptability 
to survival in dwindling waters. 

Paleohydrographic circinnstances include the 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



75 



probability that in the north-central Great Basin 
ingression of any immigrants from the relatively 
rich Columbia River fauna has been blocked by 
reason of long-standing intervening contact be- 
tween the tributaries of lakes Lahontan and Bon- 
neville. There seems to be no evidence of any 
aqueous connection by which any members of the 
Bonneville fauna ( richest in the Great Basin area ) 
could have reached any of the basins under con- 
sideration, at least in late Pleistocene time. An 
ancient connection with the Colorado River sys- 
tem, probably early Pleistocene or earlier, pre- 
sumably explains the occurrence in the Lake 
Railroad basin of Crcnichthys (pp. 35-36), but no 
other fish representative of the Colorado fauna 
.seems to have persisted in any of the endorheic 
basins under study. 

The fishes of the desert basins under treatment 
have very seldom survived in any of the very 
limited waters other than springs, .spring-fed pools, 
and .short stretches of spring outflows, and are 
very largely restricted to waters of this type that 
rise near the margins of the playas. along the 
base of the adjoining mountains, ordinarily along 
more or less active faults. Such springs, though 
less inconstant than others, sometimes, as we have 
seen, stop flowing during extreme droughts (as 
in 1934), and many of them contain relatively 
few individuals, usually so crowded as to seem 
subject to destruction by predation or disease, 
even though all surface water does not disappear. 
In some basins only one or a very few springs 
have survived. Springs that rise on mountain 
slopes, and the springs and small streams that 
flow in the mountain canyons, often give the 
illusion of constituting possible to highly propi- 
tious fish habitats, and indeed for some time — 
even for years — may support introduced fish 
(trout, minnows, or suckers), but are devoid of 
native fish for a very good reason: the occasional 
torrential precipitation characteristic of such 
desert regions, striking locally only at intervals 
of years, decades, or even centuries, transforms 
the trickling waters into raging torrents that roll 
boulders, sometimes as large as a house, and 



flush everything onto the normally dry playas. 
There, the waters usually recede and disappear 
too rapidly to permit the fish to wend their way 
backward to habitable waters. At times water may 
persist for months or even years on the playas, 
but becomes too alkaline and too ephemeral to 
support fish. Trout that succeed well under or- 
dinary conditions are occasionally, as is well 
known, wiped out, so as to require restocking. We 
have encountered several instances in which min- 
nows or trout have thus been eliminated from 
waters wherein they had persisted for a few to 
many years, even from pluvial time. 

The environment indeed has been so harsh 
that it is a wonder that any populations have 
survived. However, all four of the species that 
occur in the basins we are treating have exhibited 
exceptional ability to survive under the harsh 
conditions imposed by the very limited and iso- 
lated aquatic habitats of the arid American West. 
Rhiiuchtliys osciihis and Gila hicolor display such 
adaptation (perhaps in part preadaptation) over 
a wide area, and arc the only fish species that 
have held out in any of the Great Basin waters 
under treatment, other than the two endorheic 
.systems inhabited solely by Relictiis solitariiis 
and the one endorheic basin inhabited, along with 
subspecies of Gila liicolor, by Crcnichthys iicva- 
clac. Both Rclictiis and Crcnichthys have also 
dramatically demonstrated their ability to persist 
in isolated spring waters. In occasional stream- 
lets, especially those that maintain flow onto the 
valley floor, minnows have become, or have re- 
mained, adjusted to the tenuous conditions (for 
example. Rliinichtliys osciiliis in Indian Creek in 
Crescent Valley — p. 106). 

One circum.stance that to a degree has coun- 
teracted the depauperation of the fish fauna of 
the north-central Great Basin has been the trend 
toward raciation and even subspeciation, in cor- 
relation with the sharp isolation of the remnant 
stocks since pluvial time, not only between the 
populations in different endorheic basins, but also 
within some of these relatively small basins. Thus, 
we have found differences that we interpret as 



76 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



being on tlie subspecies level between the piipula- 
tions of Rhinichlhys osculus that have survived in 
the basins of pluvial lakes Gilbert. Diamond, and 
Clover, and between the stocks of Gila bicolor 
that have persisted in the basins of lakes Diamond. 
Newark, Clover, and Railroad. Within single 
basins we have recognized as on the subspecies 
level different popidations of Rhinichtliys osciiliis 
within the Lake Clover basin and of Gila hicolar 
within the Lake Newark and Lake Railroad 
basins. The two populations of Diamond Valley 
referred to as variants of Gila bicolor oln'sa ap- 
proach the subspecific level of separation as a 
unit and are well differentiated from one another 
racially. Slight racial distinction seems evident 
between the two stocks of Rhiniclitliyw osculuy 
oligoponis in the Clover Valley section of the 
Clover basin, and between the several populations 
of R. o. rol->ii.\tiis occupying separated springs in 
the basin of Lake Diamond. No strong differ- 
entiation, however, was noted between the now 
generally isolated populations of Rclictiis soli- 
turiiis. even between those inhabiting the two 
separated basin complexes ( Franklin-Cjale and 
Waring-Steptoe ) . Gene interflow between the 
ordinarily separated populations of this genus, 
and of the other genera, following interconnection 
dming the rare occasions of local torrential pre- 
cipitation, may well have dampened the rate of 
regional differentiation. 

LiMiPED Sympathy 

The extremely limited incidence of species 
sympatry in the aritl region under tieatment even 
more strikingly emphasizes the harshness of fish 
environments, the heavy toll by extinctions that 
must have occurred, and the resultant depaupera- 
tion of the fish fauna. Even within the three 
basins among 21 that contain more than a single 
species, the two that they harbor are sympatrie 
largely in the sense of occupying the same drain- 
age basin. In the entire area covered, of 54,028 
sq. km., encompassing a large part of one of the 
largest states in the country, two native species 



have been found, during extensive reconnaissance, 
to occur together in only three isolated springs! 
The limited degree of precise sympatry becomes 
increasingly evident when the circumstances are 
considered for each of the three basins that con- 
tain two species. 

In the immense drainage basin of pluvial Lake 
Diamond ( 8,097 sq. km., including the Lake 
Diana and Lake ^ ahoo drainage basins). Rhiiii- 
ciithys osculus occurs alone in the following 
widely separated locations: R5, Potts Ranch; 
R5A, Dianas Punch Bowl; R6, Coils Creek; and 
R8, Big Shipley Spring ffig. ?<). Gihi bicolor 
occurs alone at only one location (G6). Sulphur 
Spring. At only one place. Birch Ranch ( R7, 
G5 ), were the 2 taken together, and it is doubtful 
that they occur together elsewhere in the drainage 
basin, but here, so far as we could see. the species 
freely associate. 

In the basin of pluvial Lake Clover (fig. 12), 
of 2,624 sq. km.. Rhinichlhys osculus lives alone 
at two locations (RIO and Rl 1 ), both in Clover 
Valley, and occurs with Gila bicolor in the only 
habitable spring complex. Warm Springs, in In- 
dependence Valley (R12, Gl 1, the one spot where 
Gila lives in the drainage basin). Even in this 
small spring area the two species are not com- 
pletely sympatrie in the strictest sense, for 
Rhinichlhys is the more secretive and holds out 
in the den.ser vegetation, whereas Gila swims more 
freely in the limited available open water. They, 
therefore, though definitely sympatrie in the 
geographical sense, are not completely associated 
ecologically (they are syntopic, to use a term 
advocated by Rivas. 1964). 

Throughout the vast drainage basin of pluvial 
Lake Railroad, comprising 10,874 sq. km. (in- 
cluding the drainage basin of Little Fish Lakes), 
Gila bicolor inhabits many springs, including the 
sump basin of Lake Railroad, to which Crcnich- 
thys ucvadac is strictly confined. But here the two 
species are almost completely segregated from 
one another: Gila monopolizes the cool springs 
and Crcuichlhys takes over in the Inn springs. At 
only one place, niit in a spring proper, were both 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



77 



collected: in Duckwater Creek about three miles 
above Duckwater Store 445 specimens of Gila 
bicolor were taken on September 9, 1934, along 
with 23 strays of Crenichthys uevadae. Further- 
more, recent field work indicates that neither 
species now inhabits this creek. 

In all other drainage basins in the vast study 
area that have retained native fish only one 
species has persisted (tables 2, 3): Rhinichthys 
osculiis alone in the single fish habitat in the 
drainage basin of pluvial Lake Gilbert ( and it 
was apparently exterminated between 1938 and 
1959); Gila bicolor alone in the few fish-in- 
habited springs of the Lake Newark drainage 
basin and in all of the habitats of the species in 
the several divisions of the Lake Railroad drain- 
age complex other than the one mentioned above; 
and the relict dace. Relictiis solilarius. alone in 
the many fish-harboring springs of the Franklin- 
Gale and Waring-Steptoe drainage basins. The 
possibility that Gila bicolor occurred in Ruby Lake 
of the Lake Franklin basin prior to the establish- 
ment of bass, Micropterus salinoidcs, is, however, 
not completely excluded, though we think the 
joint occurrence to be highly improbable (p. 209). 

It would be difficult indeed to find any other 
fish-inhabited section of the world, of comparable 
size, which has been so devoid of competition be- 
tween fish species! 

Areas of Basins Correlated With Richness 
AND Diversity of Fish Faunas 

In our earlier report ( Hubbs and Miller. 
1948b) we pointed out that, in a general way, 
for the Great Basin province, a positive correla- 
tion exists between the area of a drainage basin 
and the richness of its fauna. This conclusion is 
consonant with a commonly held hypothesis for 
areas and species in general, particularly in con- 
nection with islands ( MacArthur and Wilson. 
1967, chapter 2). As is generally recognized, 
isolated drainage basins are, in effect, zoogeo- 
graphic islands. The data (tables 2-4; fig. 15) 
for the basins under present study, and for a 



major selection of other parts of the Great Basin, 
comprising in all 426,316 sq. km. (as measured 
respectively by us and by Snyder et al., 1964) 
fall in line with the general hypothesis, and seem 
reasonable on the assumptions that there have 
been many extinctions and that chances for sur- 
vival have been greater in the more numerous and 
more diverse habitats provided by the larger areas. 
The coefficient of correlation between area and 
number of species, for the total drainage basin 
of the pluvial lakes entered in table 4, is 0.56 ± 
0.10. Obviously, the diversity of the fauna, as 
indicated by the number of genera and of families 
represented (also shown in table 4), is also posi- 
tively correlated with the area of the drainage 
basin. 

The major irregularities in correlation between 
the area of drainage basins and the number and 
diversity of species, as shown in table 4, are 
largely explainable. The anomalously high num- 
ber and diversity of species for the Bonneville 
(no. I), Malheur (6), Goose (29). Truckee 
(34), Tahoe (48), Eagle (54). Long Valley 
(56), Ash Meadows (64). and Horse (67) drain- 
age basins are attributable to the long or com- 
plete connection of the waters in these basins with 
a larger water system below, having a relatively 
large and diverse fish fauna. The entries for these 
basins are all those shown on figure 15 to the left 
of and above the dashed line. Most of the larger 
basins with no fish species, or with only one or 
two, are particularly arid. 

The validity of the indicated positive correla- 
tion between the areas and the number and diver- 
sity of species is not thrown into question by the 
particular selection of basins that was made for 
the purpose of tabulation and computation. We 
have included all pluvial basins that contained a 
Pleistocene lake, as recognized by Snyder el al. 
( 1964). for the north-central part of the Great 
Basin under special study, for the Lahontan sys- 
tem, and for the area farther northwest, and for 
the Death Valley system (Miller, 1946. 1948): 
also for Lake Bonneville. We do not include "Hot 
Creek Lake" and the "lake in Esan Valley." 



78 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 4. Corrclaticn of lucas of sclct U-d Great Busm pluvidl-lakc draiim-^f hus'uis with rufmcss and diversity oj 
native fish fauna. 

ItKliidiny all basins tliat contained a I'lfislotcne lake, as lisled by Snyder et ul., wiili evceptions discussed in tlie text (p. 77). Lakes with names 
marked by an asterisk are those dealt with in the present report. Data graphed in figure 15. 





Pluvial (ir 


ainage basin 






Pluvial di 


ainage basin 








Area 


Fish 






Area 


Fish 


1 


Niinie of lake 


sq. km.' 


fauna' 




Name of lake 


sq. km." 


fauna- 


1. 


Bonneville 


13S,112 


4-9-21 


38. 


Mono ( Russell ) 


2,056 


0-0-0 


-> 


Lahontan 


I0y,614 


4-9-11 


39. 


Alkali 


2.038 


1-1-1 


3. 


Manly 


.'i3.224' 


3-5-11 


40. 


*Gale 


1.933 


1-1-1 


4. 


Panamint 


20.629 


3-4-4 


41. 


Meinzer 


1,922 


1-1-1 


5, 


Scarles 


16,408 


3-4-4 


42. 


■ Pme ( Wah Wah 1 


1.912 


0-0-0 


6. 


Malheiii 


14.134 


4-S-9 


43. 


Haiper 


1.865 


0-0-0 


7. 


Owens 


10,969 


3-4-4 


44. 


High Roek 


1.813 


1-1-1 


S. 


Railroad 


10,874 


">.''.'> 


45. 


Hubbs 


1,677 


0-0-0 


4. 


Tecopa 


10, -"567 


2-3-5 


46. 


Indian Springs 


1,660 


0-0-0 


10. 


Meihave 


10.502 


l-l-l 


47. 


Gilbert 


1.528 


1-1-1 


II. 


Wariny 


9.41 1 


1-1-1 


48. 


lahoe 


1.427 


4-7-9 


12. 


Mani\ 


9,363 


1-1-1 


44, 


Diana 


1.339 


0-0-0 


l.v 


Diamond 


8.097 


1-2-2 


50. 


1 unai Crater 


1.321 


0-0-0 


14. 


C atlou (incl. Guano 1 


7,656 


1-1-1 


51. 


liuffalo 


1.3 18 


0-0-0 


l.S. 


Warner 


6,S58 


2-4-4 


52. 


C arpenter 


1.257 


0-0-0 


16. 


Dixie 


6.314 


1-1-1 


53. 


little Fish lakes 


1,131 


1-1-1 


17. 


Alvord 


6,066 


-»_-).-> 


54. 


Fagle 


1.127 


3-5-5 


IS. 


*Steptoe 


5,462 


1-1-1 


55. 


Jake 


1,114 


0-0-0 


ly. 


■Franklin 


5,271 


1-1-1 


56. 


Long Valley 


947 


2-3-3 


20. 


Fort Roek 


5,219 


2-3-3 


57. 


Delamar 


997 


0-0-0 


21. 


• Spring 


4.337 


()-(l-(l 


58. 


C'arico 


966 


1-1-1 


">"> 


Crescent 


4,237 


1-1-1 


59. 


Cave 


422 


0-0-0 


23. 


C'hewancan 


.^.859 


2-3-3 


60. 


Kumiva 


917 


0-0-0 


24. 


Surprise 


3,836 


''.''.'> 


61. 


Antelope 


870 


0-0-0 


2.'i. 


■ Newark 


3.587 


1-1-1 


62. 


LIppei Lake Stepto 


e 816 


0-0-0 


26. 


Toiyahe 


3,354 


1-2-2 


63. 


Lahou 


777 


0-0-0 


27. 


Bristol 


2,979 


ll-(l-0 


64. 


Ash Meadows 


756 


2-3-4 


2,S. 


Adobe 


2.901 


0-0-0 


65. 


Washoe 


251 


1-1-1 


2y. 


Goose 


2.875 


5-7-7 


66. 


Lemnion 


241 


O-O-O 


30. 


Fish (White Mts. ) 


2.779 


1-1-1 


67, 


Horse 


210 


"'.'».'> 


31. 


* Clover 


2.624 


1-2-2 


68. 


Spanish Spring 


186 


0-0-0 


32. 


Pahrtimp 


2.569 


1-1-1 


64. 


Sn\der 


148 


0-0-0 


33. 


Coal 


2,541 


0-0-0 


70. 


Laughton 


124 


0-0-0 


34. 


Truckee 


2.479 


4-7-8 


71. 


Summit 


48 


0-0-0 


3.^. 


Granite Springs 


2,476 


0-0-0 


72. 


Fred 


57 


0-0-0 


36. 


Wellington 


2,383 


0-0-0 


73. 


Yahoo 


44 


0-0-0 


37. 


Madeline 


2.163 


1-1-1 











1 EnUiL- arcii tributary to lake nanicd, iiKUitlinj^ areas (also tallied) of basins of lake within the total watershed. 

- Families — genera — species. 

'Including Panamint, Searles, and Owens (excluded by Snyder et tiL). 



wliich wc do not accept, at least a.s late Pleisto- 
cene, nor Lake Reveille, which we interpret as 
having been part of Lake Railroad at niaxiinum 
level. The drainage basins that we have omitted, 
among the total pluvial-lake basins as listed for 
the Great Basin by Snyder et al. ( 1964). are all 
wholly fishless, and they are all in the southern 
part of the Great Basin, south of the Lahontan 
system and the basins particularly included in 



the present study, in an area that is now extremely 
arid and was presumably moderately arid even 
during the height of the pluvial periods, as we 
have stressed (Hubbs and Miller, 1948b), Many 
were prc^bably shallow or even intermittent or 
dubious, and not one appears to have left a bold 
shoreline. The .'^S fishless drainage basins in- 
volved, as listed, contained 45 ]iliivial lakes, of 
which Lake Columbus is thotmht to have re- 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



79 



23 



CO 
LJ 

O 
LJ 

a. 

CO 



Li_ 

o 

LiJ 
DD 



18 



13 



CORRELATION COEFFICIENT r = 0.56±0I0 



J L_ 



/ 
/ 
/ 
/ 



+6 / 
/ 
/ 
/ 



,15 7 ,5,1 



,55 / 



26 

3IA24 J 7 13, 



m il I II ++HH-* 



+ +-»-+ + ++ 

21 
+ 
I 



Figure 
species. D 



1 2 3 4 5 6 7 8 9 lOII 12 

NATURAL LOG., AREA OF DRAINAGE BASINS OF PLUVIAL LAKES IN SO KM 

15. Correkition hctwcen ;uca of selected pluvial-lake basins in (ireal Basin and niimher of nati\e fish 
ata and lake nunihers from table 4. 



ceiveci the outflow of Lake White Mountains 
( Fish ). which has a single native fish species, and 
is included in table 4. The listed aieas of these 
38 basins range from 78 to 8,094 sq. km.; only 
11 exceed 2.000 sq. km. 

Scrutiny of the data for the preceding discus- 
sion of the relation between the area of basins and 
the richness and diversity of the fish faunas 
throughout the Great Basin indicates that the 
north-central part, which we are particularly treat- 
ing, is representative of the whole area, except 
for the three largest and faunally richest basins, 
those of lakes Bonneville, Lahontan. and Manly. 
The key feature is the trend toward fish-faunal 



extinction in the smaller, more isolated, and more 
arid drainage basins. 



Char.\C'ters and Systematic Status 
OF Species 

Our systematic analyses of Rliinicluhys osciiliis. 
Gila hicolcr. and Rcliclu.s sclittiriiis. of the sub- 
species and local races of the first two, and of the 
local populations of the relict dace, have been 
based, primarily but not exclusively, on various 
measurable and countable features. Various items 
of size, general form, coloration, development of 
sensory structures (pores and barbels), dentition. 



80 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



dinmiphism in secondary sexual characters, num- 
bers and size of individuals, and bioniass are 
accorded attention. Other emergent methods of 
evaluating relationships, such as biochemical tests. 
karyotype analyses, behavior studies, and genetic 
experiments, would no doubt add nuich to the 
security of conclusions and to a fuller under- 
standing of the origin and differentiation of the 
various isolated populations. Except for a start 
on karyotyping (p. 19.^). these promising ap- 
proaches almost wholly remain to be undertaken. 
Tran.sference experiments and the analysis of 
characters of fishes incidentally established into 
different types of habitat may often be a propi- 
tious means, and the most feasible way. to infer 
the genetic fixity of characters or the direct effects 
of altered environments ( indeed one such test — 
in effect a natural experiment — was made by us 
on a population of Rhiiiichthys osciiliis that had 
been transferred, as bait minnows', from one of 
the headwaters of Humboldt River '\n{o an iso- 
lated spring in Ruby Valley — p. 107). It is our 
hope that our studies will encourage others to 
e\pand the knowledge of the remnant fish life of 
the arid American West. 

The characters of the fishes in the north-central 
Great Basin very largely reflect their adaptations 
to the spring habitats to which they are mainly 
restricted. These features, which they tend to 
share in common, are treated below, under the 
heading of Color, Texture. Form, and Sensory 
Structures. 

The sharply restricted and concentrated ranges 
of the forms under treatment, along with their 
abundant populations, have made it possible to 
obtain comprehensive material for an analysis of 
the patterns of variation in various characters, 
and to learn something of population structure. 
It is indeed seldom that samples can be obtained 
so representative of whole populations through- 
out their range. We have tried to take some 
advantage of this potential, but fiuther studies 
are surely in order. 

We now proceed to a discussion of the several 
sets of characters that we have analyzed, treating 



definitions, methods, and procedures, as well as 
offering some evaluation of the characters. The 
actual presentation of the basic data, and their 
application to specific problems, are deferred to 
the accounts of each form or population. The 
order of presentation in the following discussion 
mainly follows the sequence adopted in the de- 
scriptions of the various local forms. 

Size. 

For all collections studied, we present the 
range in .standard length for all specimens and 
indicate whether the fish are dwarfed, as are 
many isolated forms of restricted waters. It de- 
velops that the members of some of the isolated 
populations of greatly restricted range herein 
treated instead of being markedly dwarfed are 
larger than usual. For example, in Rliinichfhys 
osciilus. although R. o. Icthoporus of Warm 
Springs in Independence Valley and the popula- 
tion of atypical R. o. rohiisiii.s of the warm springs 
of Potts Ranch and the nearby Dianas Punch 
Bowl in Monitor Valley are greatly dwarfed, R. 
o. icliqiiiis of the Grass Valley spring, the Coils 
Creek population of R. o. robustus, and especially 
the Indian Creek stock of that sub.species. are un- 
usually large for the species. Gila hicolor isolata 
of Warm Springs in Independence Valley is some- 
what dwarfed, whereas G. h. euchila of Fish 
Creek Springs in Newark Valley is unusually large 
and massive for an isolated population. The dif- 
ferent populations of Rdictiis solitariits also vary 
widely in the si^e attained: the largest specimen 
among each of 21 collections containing 147 to 
1.073 individuals had attained the standard length 
of SI to 94 mm., but one other lot (Collection 
17) included many large specimens, up to 114 
mm.; 7 lots, in contrast, represent apparently 
dwarfed stocks, with greatest standard lengths of 
41 to 77 mm. Very large specimens, incidentally, 
tend to have carried growth changes in propor- 
tions to an extreme ( p. 2 1 4, fig. 51). As a whole, 
however, the native fishes of the isolated waters, 
many of greatly reduced area, are small. 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



81 



Color, Texture, Form, and Sensory Struc- 
tures. 

The remnant fishes of the north-central Great 
Basin, as of other parts of the arid West, exhibit 
marked convergence in this complex of characters 
by reason of their long restriction largely to small 
isolated bodies of water with little current and 
with little or no competition and predation. They 
tend to be relatively sluggish, seeking shelter close 
at hand, rather than by dashing far away ( as is 
noted below). They are in general midwater 
swimmers in quiet water. Seemingly molded by 
adaptation to their physical and biological en- 
vironment, they tend to share various characters, 
which have mostly been pointed out already 
(Hubbs, 1940, p. 201; 1941a, p. 68; 1941b, p. 
1 87 ; Hubbs and Miller, 1 948b, pp. 5 1 -52). These 
features are notably exhibited by the highly re- 
stricted spring dace Erciniclilhys acros (Hubbs 
and Miller, 1948a, pp. 20-27, pi. 1 ). 

These spring fishes tend to be small, or even 
dwarfed; to be plainly and dully colored; and 
they, especially the females, tend to be relatively 
soft in flesh and texture. In correlation with life 
at mid-levels in quiet water, they tend to be 
chubby in general form, with the contours of head 
and body well rounded; and the ventral contour 
ordinarily approaches the dorsal contour in curva- 
ture, rather than being flattened as it is in related 
fishes adapted to current. The spring fishes are 
rather poorly streamlined, with a relatively deep 
caudal peduncle ( not slender and with expanded 
procurrent caudal rays as in swift-water forms ) . 
Their mouths are usually more or less enlarged, 
terminal, and oblique, and are thus fitted for mid- 
water feeding (not small, inferior, and horizontal, 
as in bottom-feeding rheophiles). Their fins, 
seemingly ample for limited locomotion in quiet, 
restricted waters, tend to be small, short, rounded, 
and supported by rays reduced in number and 
strength ( rather than being elevated or prolonged, 
falcate, and with rays strong and not reduced in 
number, as in rheophiles). Their anterior parts 
are generally enlarged, at the expense of the often 
considerably shrunken posterior parts; as a con- 



sequence the dorsal and anal fins are commonly 
set far back. The scales of the spring types tend 
to be loosely arranged, poorly imbricated, deeply 
embedded, and often with the radii on all fields. 
Particularly notable is the tendency toward reduc- 
tion or even obsolescence of the sensory canals 
and pores, the barbels, and other dermal sense 
organs, on both body and head; seemingly such 
structures are not very critically needed in the 
reduction or absence of predation. 

The extent to which these trends have been 
carried in the spring-inhabiting fishes of the Great 
Basin is outlined in the descriptive accounts of the 
various forms. Such degeneration is particularly 
well illustrated by the isolated forms of Rhinich- 
tliys osciilus and is moderately evident in Relictiis 
(p. 181). Tendencies in that direction are shown 
by the isolated races of GiUi bicolor. 

Development of barbels. The degeneration 
of the barbel is particularly striking in the iso- 
lated forms of Rhinichthys osciilus. the only 
species under treatment that ever develops this 
sensory structure. The degree of development of 
the barbel is discu.ssed under the heading for that 
minnow (pp. 100, 104 ), and, in quantified detail, 
in the accounts of the respective subspecies. 

Development of lateral-line pores. The 
tendency for the lateral-line pores to degenerate 
in the isolated spring populations involves both 
those along the lateral line of the body and those 
on the head canals. Some complexities arise in 
quantifying the degree of development. The mea- 
sure chosen is the number of pores, which need 
to be separately tallied by size groups of fish, be- 
cause the pores are not formed in the very young 
and develop rather slowly, increasing even into 
half-grown stages ( see figs. 25 and 26 and table 
12 for Rhinichthys). 

The first pore counted ordinarily lies well be- 
hind the pore at the junction of the supratemporal 
head canal with the main lateral line. The ar- 
rangement, however, is subject to potentially con- 
fusing variation. When, not infrequently, no pore 
opens at the junction, the pore following, usually 
very closely, is disregarded in the count. The 



82 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



pore tliat is usually counted first foinis very early, 
so that many young specimens yield a count of 1. 
Near the beginning of the lateral line on the bcidy 
numerous aberrancies in the canal and pores oc- 
casionally occLU'. such abnormalities as breaks 
and branches and the development of two pores 
on a scale. Especially in forms with a reduced 
lateral line, there are many irregularities along 
the whole line, with very fret]uent interruptions. 
Development of pores usually iirogresses from 
fore to aft, but as the formation ]irtKeeds. a sec- 
ondary center often appears posteriorly, and al- 
most always this occurs well before the caudal 
base. In larger specimens, however, pores some- 
times form on the terminal scales, including oc- 
casionally line or two scales over the caudal-fin 
base ( such pores have been included in the 
count ). Interruptions, often extremely numerous, 
cause complications, for isolated single scales 
often form a tube with a pore at either end. Pores 
often form on either side of a break that is later 
closed in, so that only one is now counted where 
two would have been enumerated earlier. For 
such reasons, degenerative tendencies may lead to 
increa.scd counts. In this study we have held to 
the criterion of counting each separate pore, even 
though the roofing-over of the canal is not com- 
plete. Often pores can not be detected where 
scales have been removed: for this reason, some 
counts are doubtless somewhat too low. How- 
ever, care was taken to count specimens with few 
lost scales, and such omissions as have occurred 
have presumably not seriously vitiated the 
presentations or conclusions. 

The lateral-line canals and pores of the head 
of the cyprinid species were extensively examined, 
and were found in general to follow the usual pat- 
tern for American genera as outlined by lllick 
(1956). For all forms treated, the degree of 
interruption of the supratemporal canal seems to 
be an important feature, and was tallied, with 
some counts: ordinarily the interruption occurs 
on the midline, and can be determined under due 
magnification with the use of a jet of compressed 



air. For Rcllcins soHftiriiis. counts are presented 
of the preoperculomandibular pores. 

These sen.sory characteristics are more closely 
correlated with the environment than with rela- 
tionship, as is noted above. 

In correlation with their reduced sensory equip- 
ment, the spring-inhabiting remnants exhibit less 
escape reaction than stream-inhabiting relatives. 
Thus, the creek race of Rhinlilillnw osciiliis in 
Crescent Valley showed much more escape be- 
havior than the spring form in Crass Valley (p. 
1 06 ) . A special adaptation of spring forms seems 
to be their propensity to remain in the safety of 
the spring heads, avoiding impermanent outlet 
ditches, as noted for Giln hicolor in Sulphur 
Spring. Location G5. 

Sc At i; Struciiirh. 

Scale structure (fig. 46) was examined and is 
briefly described for the forms of Rhliiichthys 
oscii/iiy and (iiUi hicolor. and for Rcllcliis soli- 
tariiis. with particular reference to the occurrence 
or lack of radii on the lateral and posterior fields 
as well as the anterior field: to the sharpness of 
the distinction between the fields; and to the 
general shape, whether essentially oval, or shield- 
shaped with strong angulation of the circuli along 
the line between the posterior and lateral fields, 
in this last respect considerable variation was 
found in (JiUi hicolor, but a preliminary suggestion 
of regional differentiation was not confirmed by 
further exanunation. 

MURIMIOMI I KV. 

In measuring the various bc^dv and heael parts 
to check on local variation in the proportions we 
have adhered to the following methods (essentially 
as proposed by Hubbs and Lagler. 1964. pp. 24- 
26). Precision dial calipers were used and read 
to 0.1 mm. and all measming was from point to 
point, by one of us (C.L.FI.). usually on consecu- 
tive days, to help insure uniformity. Appropriate 
magnification and illumination were employed. 

Our mimerous samples of the Great Basin na- 
ti\'e fishes proved favcrable for proportional mea- 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



83 



suremeiits. because they were uniformly hardened 
in 10 percent buffered formahn, then transferred 
in due time, after thorough washing, into ethyl 
alcohol. In general, the fish preserved well, with 
little twisting or opisthotonus. Furthermore, the 
skin remained firm and leathery, and the fins 
were tough, so that the specimens were little 
damaged (except for fraying of the caudal in 
some specimens), even when they had been 
carried (regularly in full glass jars) on the rough 
roads of 1934 and 1938. For most collections, 
the large series helped to prevent damage in 
transit and rendered it feasible to select for mea- 
suring very well preserved specimens of both 
sexes, of graded size. 

As a reasonable check on differences in body, 
head, and fin parts the following measurements 
were taken: 

1 . Standard length in mm. (consistently used, 

even when not so stated). 

2. Predorsal length (one tip of calipers was 

inserted at base of first rudimentary 
ray). 

3. Anal to caudal (from extreme front of 

base of first anal ray to end of hypural 
on midline). 

4. Pelvic to anal (from insertion of pelvic 

fin to origin of anal fin, as in item 3 ). 
Taken only for Gila hicolor. which 
shows some difference between pop- 
ulations in this dimension. 

5. Body depth (greatest ). 

6. Peduncle depth ( least depth of caudal 

peduncle, disregarding a slight con- 
striction, when it occurred near origin 
of procurrent rays). 

7. Head length (including opercular mem- 

brane ) . 

8. Head depth (occiput to isthmus, even if 

slightly oblique). 

9. Head width ( greatest ) . 

10. Snout length. 

11. Orbit length (may be oblique: between 

true fleshy rims of orbit, disregarding 
puffy conjunctival tissue). 



1 2. Upper-jaw length ( to end of maxillary 

under suborbital). 

13. Mandible length (omitted for Rhinicli- 

thys. in which the mandible is small 
and the mandibular joint is difficult to 
perceive ) . 

14. Intcrorbital width (least). 

15. Suborbital width (very definitive, by tak- 

ing care to avoid puffy conjunctival 
tissue). 

16. Depressed dorsal (from structural base at 

front, where one point oi calipers was 
appressed rather firmly, to extreme tip 
of whatever dorsal ray extended far- 
thest). The anal fin was not measured, 
as its length seems to be correlated very 
closely with that of the dorsal fin. 

17. Caudal-fin length (from base of middle 

rays to extreme tip of longest ray, in 
either lobe; since caudal rays are partic- 
ularly subject to breakage, fewer mea- 
surements are often given. Care was 
taken to limit this measurement to 
specimens with at least very nearly 
complete ray tips). 

18. Pectoral-fin length (extreme). 

19. Pelvic-fin length (extreme). In measuring 

the paired fins (from upper/outer 
base ) , care was taken to avoid the often 
commonly broken tips, and often the 
larger of the paired fins was measured. 

In addition, the longest gill-raker was mea- 
sured, in GiUi hicolor only, as discussed below ( p. 
146). 

The measurements, transformed into permil- 
lages of the standard length, are presented sepa- 
rately by subspecies or populations for large-size 
groupings of each sex. Further analysis of 
Rhinichthys osciilus and Gila hicolor is presented 
for only a few, more critical dimensions. More 
detailed morphometry was carried out for popula- 
tions of Rclictiis soUtariiis. The large mass of 
primary measurements have been retained and 
can be made available should occasion arise for 
further analysis. 



84 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



The liniilcd analysis so far atteniptod shows not 
only some useful differences between populations, 
but also demonstrates changes with age and the 
rather sharp sexual dimorphism that develops in 
all three cyiirinid genera in the area. 

Si:.\UAi Dimorphism and Nuptial CirxRAcriRs. 

In all three species under detailed treatment, 
as in most cyprinids. males contrast with females 
in their smaller size, appreciably firmer flesh, and 
less posteriorly inserted dorsal fin. In other 
respects, the fins exhibit conspicLious sexual di- 
moiphisni. becoming in the males not only rela- 
tively longer but also stronger, with thicker rays. 
The pectorals and the pelvics become strongly 
modified in the nuptial males, with the anterior 
lor outer) rays and surrounding tissues especially 
enlarged; and these fins become more expanded. 
The pectoral in particular becomes greatly dilated 
transversely, stands out. and is somewhat twi.sted. 
The elistinction is developed early, and increases 
with age and size, except that some very large 
females were noted to slightly approach the condi- 
tion of males. 

The nuptial tubercles, especially on the pectoral 
fin, are sharply distinctive in the three genera. 
Those on the head are outstandingly diagnostic in 
Rclictii.\. 

Sexual dimorphism and nuptial tubercles are 
treated under the headings of Rliiiiicliilns o.sriiliis 
( p. 98 ) and subspecies (pp. 113. 121. 125. I 3.^ ) ; 
subspecies of Gila hicclor I pp. 152. 15(1. 161. 
166. 172. \19): [he genus Rclicliis (p. 1X2) and 
the species R. solilariiis (p. 219). 

Mrristics. 

Particular emphasis was placed on meristics. 
as the numerical characters are in general pre- 
cisely determinable, free of changes with age, and 
readily subject to analysis. Meristics provide some 
of the most striking differences between the iso- 
lated populations. 

Furthermore, the abundance of material of 
relatively uniform genetic constitution has offered 
a favorable opportunity to determine the extent 



of variability in the meristics of some structures, 
particularly of fin-ray and gill-raker numbers. The 
extent of variation determined is surprising, and 
raises the question that we do not solve, but do 
present data for solution, as to whether such fac- 
tors as limited predation and inbreeding in the 
highly localized populations may have led to in- 
creased variability. The unexpected degree of 
variation in the number of caudal rays (p. 86), 
for example, opens up this problem. 

Moreover, the presumed and apparent genetic 
integrity of the highly localized populations, some 
inhabiting a single spring, provide unusually 
fortunate material for testing some of the basic 
interrelatiimships between the numbers developed 
in different structures, for example: the cor- 
relations between the numbers of dorsal and anal 
rays, between the numbers of pectoral and pelvic 
rays, and between the numbers of left and right 
pectoral and pelvic rays. A special point is the 
determination in such propitious material of the 
degree of asymmetry and the degree of dextrality 
in the number of rays in the left and right paired 
fins, along the lines developed by Hiibbs and 
Hubbs ( 1945). Data on such relations are pre- 
sented in the following discussion of the meristics 
of different stitictures. 

A shadow of uncertainty is always inherent in 
the use of meristic characters in taxonomy, and 
in variational analysis, by the circimistance that 
some of the variation in the numbers oi serial 
parts may be the direct effect of the environment. 
This is one of the cogent reasons why an experi- 
mental approach should be added in the study of 
such pi\iblems as we are now treating. Since en- 
vironmental temperatures are a major factor 
known to affect the serial numbers, we have made 
one approach, albeit rather crude, to evaluating 
whether the observed differences may be geneti- 
cally fixed by correlating the i.letermined mean 
values with the temperatures found in the habi- 
tats — a procedure that is especially applicable in 
spring waters, which tend to be unusually constant 
in temperalurc. No obvious correlation was noted 
for any of the 3 cyprinid species studied, nor was 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



85 



Table 5. Correlation between mean number of fin rays and of vertebrae (V) and observed summer water tempera- 
ture of habitats sampled for populations of Rhinichthys osculus in certain basins in Nevada.^ 



Location 


'a 


D 


A 


P, 


P= 


V,, 


V, 


V, 


Coils Creek (R6) 


13.9 


1.11 


7.00 


13.64 


7.79 


19.85 


17.44 


37.50 


Crescent Valley (Rl ) 


17.2 


7.88 


7.00 


13.84 


7.97 


19.76 


18.14 


37.90 


9.5 mi. S. of Wells <Rin» 


17.2-17.8 


111 


6.98 


12.49 


6.95 


20.31 


16.69 


37.00 


Grass Valley (R9l 


18.9 


8.09 


7.05 


12.65 


7.18 


19.97 


16.51 


36.44 


WarmSpr., Clover V. (RID 


19.3 


7.93 


7.00 


12.72 


6.96 


20.05 


17.37 


37.55 


Carico L. Valley ( RO ) 


20.6 


8.0? 


7.0? 


12.3? 


7.8? 


20.3? 


18.0? 


38.5? 


Bishop Creek (R2) 


21.1-22.2 


7.97 


7.00 


13.83 


7.88 


18.75 


18.06 


36.76 


Town Creek ( R3 ) 


21.7 


7.96 


6.93 


13.65 


7.68 


19.35 


18.23 


37.54 


Birch Ranch (R7) 


22.2 


7.86 


6.97 


13.31 


7.22 


18.73 


17.75 


36.63 


Independence V. (R12) 


25.6 


7.94 


6.95 


12.72 


6.97 


19.45 


17.24 


36.68 


Big Shipley Spr. (R8) 


27.8 


1.11 


6.86 


13.29 


7.23 


19.56 


17.84 


37.42 


Potts Ranch ( R5 ) 


31.7-34.4 


8.00 


6.98 


13.35 


7.68 


19.26 


17.18 


36.59 


Dianas Punch Bowl ( R5A ) 


37-39.0 


8.02 


7.04 


13.67 


7.63 


19.00 


18.35 


37.50 



1 Data graphed as figures 16 and 17. 

- In the accounts of the Locations the temperatures have been rounded to the nearest degree. The temperatures for the three lots from Diana 
Puncli Bowl Sprmgs were taken in Fehniarv. and the data for this locality are not graphed in figures 16 and 17. 



any indicated by analysis of the available data for 
the vertebrae and fin rays of the several subspecies 
and races of Rhiitichthys (table 5: figs. 16. 17). 
Fin rays. The rays were counted under ap- 
propriate magnification and illumination, accord- 



























<-> 


1 


5 
E 


o5 


1" t 5 

QD t— m 


4; 


0^ 


c 
a 
CE 

1 




20 
19 
18 


i 




-^ 


T f 


.ti 


i 


» 


— ' — 


_l UJ 

< < 
Q q: 

3 CD 


• 




• 


• 


•• 


■ 


• 




UJ LC 

cr UJ 
























19 


















UJ 


















<^ 


18 


- 




• 




. — • 








^^ 


17 


' 




__ 


* 




■ 






^ UJ 

> 




"^Q 


/I6 




































UJ 

< 


38 


\ 




• 


. 


■ 








\~ UJ 

PS 

LU 
> 


37 


^v-J 






• 




I 


1 




I 



15 20 25 30 

OBSERVED WATER TEMPERATURE, »C 



35 



Figure 16. Apparent lack of any definite correlation 
between mean number of vertebrae and recorded water 
temperatures for 12 populations of Rhinichthys osculus 
from basins in Nevada. Data specified in table 5. 



ing to the methods standardized by Hubbs and 
Lagler ( 1964. pp. 19-21). 

As noted below, in the discussion of vertebral 
numbers, there seems to be no significant correla- 
tion between the number of fin rays and the tem- 
perature of the spring-water habitats (fig. 17), 
strongly suggesting that ray numbers are not de- 
termined by the ambient temperatures. 

Dorsal and anal rays. The count of dorsal 
and anal rays is of principal rays, and no compli- 
cation arose as to the first ray to be enumerated, 
which was always unbranched and essentially of 
full length. The last two .serial elements were con- 
sistently counted as one ray, even though, oc- 
casionally, they were largely conjoined, or the 
last element was a minute rudiment. Such condi- 
tions are especially frequent when the ray number 
is higher than the norm. Conversely, when the 
number is low. the second element of the last ray 
is often so far removed from the first unit as to 
simulate what might be interpreted as a non- 
doubled last ray. 

The number of dorsal rays (tables 15. 29. 41 ) 
is sharply modal at 8 in all populations of all 3 
species under detailed treatment, as it is in a 
large proportion of small American cyprinids. 
The observed variation is only 7-9 in Rhinichthys 
osciihis and 7-10. with counts of 9 and 10 ex- 
tremely rare, in Gila hicolor and in Relictiis 



86 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 











_, 


















'V 
































*■ 




"G 


■— -^ £ 


^ 


3^ 


j= 








































" 






>Lrt 


a. o Q. 


-0 

c 




cc 




o 


t 


t 
in 




S ° c ^ 


15 


^5. 


1 




* 




- 


44 




* 


* 


~ 




- 




, 


• 


: 


, 






8 






• 




' — " 












7 5 




70 


-• 




-- 




■ ■ ^ f 


• 


• 




140 


- 




• 












135 










• 




• 




130 
















• 






12 5 


- 




■ — • 




! 








80 


- 




• 




• 








7 5 




7 


1 


! 




■ 


1 


1 


1 





15 20 25 30 "- 

OBSERVED WATER TEMPERATURE. °C 

Fu.UKL 17. Apparent lack of an\ dclinilc corrcla- 
lion helween mean niiniher o\ I in ra\s and recorded 
sLininier water temperatures lor 12 populations of 
Rliitiii htin s iiMiilus from hasnis in Nevada. Data 
specified in table .*>. The receniK ohtamed supplemen- 
tary data for Dianas Punch Bowl Springs, lor recordctl 
temperatures of .^V-.^"-'.!) C.. deviate from those for 
Potts Ranch as follows; dorsal. +.02; anal, +.0h; 
pectoral. !..''2; pelvic, -.05. 

solilariiis. The occasional variants are more 
often down than iip in Rliiiiivluliys and Rclicliis. 
but this is not consistently true for Gilii and is not 
Irue for Rh'm'ulithys oscuhis rclic/iiiis. 

The anal rays (same tables) are sharply modal 
at 7 in all the populations of Rhinichthys osciiliis 
and Rclictus soUtarhis. both of which usually in- 
habit springs and small creeks, but deviate in the 
Gila hicolor populations from the more usual 8 



TABt.E h. C'dinldlion hctwft'n niiDihcis of dorsal iind 
(inal rays in selected samples of exprinid fislies in llie 
north-central Great Basin. 



No. of No. of anal rays 

dorsal 

rays 5 6 7 8 



R/iinichlhys osciilus lohiistiis: 1 

5 Locations. 1.. Diamond system S 

9 

Rhuiichlliys osciilu\ itliiiuiis: 1 

I . tiilbcrt s\sicm 8 

9 

(j'//i/ hicolor ncwaikcnsis: 1 

3 Locations. L. Newark s\stem 8 

9 

Rclictus solitariiis ; 7 

6 Collections, whole range 8 

9 
10 



1 



5 20 — 

4 223 4 

— 24 

2 

5 65 6 

— 7 3 

— 11 — 
2 117 14 

— 7 — 



I 



7 — 

5 98 2 

— 1 — 

— 1 — 



toward or to 7 in the .i subspecies restricted to 
springs. Thus the trend toward reduction in counts 
in spring-inhabiting populations is illustrated. The 
ob.served ranges are 5-<S for Rliiiilchlhys osciilii.s, 
6-9 for Gild hicolor. and b-H for Rclictii.\ .soli- 
tariiis. 

The numbers of dorsal and anal rays were 
found to be es.sentially independent characters, for 
the positive correlation between these values is at 
most very low (table 6). as is usual for dorsally 
and ventrally located structures that are not di- 
rectly opposite. 

Caudal ra'is. Even though the caudal rays 
seem to vary extraordinarily in number in all 
three species deah with (table 7 ). the outermost- 
marginal of the principal rays (the ones counted) 
were always extra-wide and unforked. Occasional 
lack of branching of an inner ray was disregarded. 
Seemingly interpolated, generally weaker than 
average rays were counted, even though isolated 
m the fin membrane (but these did not give rise 
to the counts of 20 or 21). Occasionally two 
caudal rays were nu^re or less fused, yet were 
counted separately, to give, ordinarily, the modal 
number ( \9 ) . 

The caudal rays in all three species are sharply 
modal at 1 9. as expected, for this is the number that 
is chatacteristic of the family Cyprinidae through- 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



87 



Table 7. Numher of caudal rays in all populaiinns 
of each native cypiinid fish treated.^ 

Number of principal caudal rays 
14 15 16 17 18 19 20 21 No. Mean= 



Rhinichlhys 
<> senilis 1 

Gila 

tiicolor — 

Reliclus 
solitarius — 



1 5 49 450 25 4 535 IS.'i.^ 
1 3 22 329 12 — 367 18.95 
1 3 8 82 8 2 104 18.95 



iData from tables 15, 29, 41. 
- Mean for summarized counls. 



out its range, and of most other nialacopterygian 
fishes. Because of this consistency, and because 
the relatively recent trend to count caudal rays 
is showing that the number of rays in normally 
forked caudal fins is generally constant within 
whole families of fishes, it is amazing that a wide 
range of individual variation, remarkably similar 
in pattern, has been encountered in all three 
species ( table 7 ) : 1 6-2 1 , with one aberrant 
count of 14. in Rhinichlhys oscuhis: 16-20 in 
Gilii hicolor\ and 16-21 in Rchctiis soUtaiius. 
This wide fluctuation is wholly age-independent, 
and does not seem to be population-dependent, 
for the modality at 1 9 is so strong and constant, 
and nearly all of the individual populations dis- 
play variation in the number (tables 15, 29, 41 ). 
Maximum observed ranges for single samples are 
5 rays in one sample, and 4 rays in 5, of the 13 
samples, in Rliinichthys: 5 rays in 1 sample and .""^ 
rays in 5 samples (out of I I ) in Gila; and 4 rays 
in 5 of the 6 categories for Reliclus. There seems 
to be a greater tendency to decrease than to in- 
crease the count. The decrease can occur within 
a caudal lobe. One Rhinichlhys specimen had the 
third and fifth rays from the lowermost so ex- 
tremely slender as to be detectable with difficulty. 
The mean number of rays is almost identical in 
the three species. It is queried above whether this 
rampant individual variation in a character nor- 
mally constant may be related to the population 
structure of these isolated-spring fishes. There is 
no indication that any of the spring-limited forms 
exhibits an increased trend toward a decrease in 



caudal-ray number, despite a weakness of the fin. 
However, perhaps in correlation with life in very 
restricted springs, a much greater reduction has 
been found to characterize a notably dwarfed 
form of the Rhinichlhys osciilns complex that lives 
in very small springs in Snake Valley in the Bon- 
neville basin ( immediately east of the area under 
treatment). 

Paired-fin rays. Pectoral and pelvic rays 
were all counted, without regard to branching or 
size, even though the lowermost (innermost) ray 
was so extremely minute that strong magnification 
and optimal lighting were required for its percep- 
tion. 

The pectoral rays fluctuate widely in number 
in all three species, probably in correlation with 
the moderately high number of rays and the 
sharp reduction, often to obsolescence, of the rays 
toward the lower margin of the fin. In any one 
of the populations herein treated, the usual range 
is 2-4 rays, and the total range observed within 
a species is 8 rays (9-16. very rarely fewer than 
12) in Rhinichlhys osctiliis and in Rclictiis soli- 
lariiis, and 7 rays ( 13-19) in GiUi bicolor ( tables 
15. 30, 41 ). There is no consistent interpopula- 
tion variance in the populations .studied of either 
Gila or Reliclus. 

The three recognized isolated endemic sub- 
species of Rhinichlhys oscuhis ( table 1 5, fig. I S ) 
have a somewhat reduced average number of 
pectoral rays (12.49-12.72 vs. 13.29-13.89 in 
R. (>. rohusius with an exceptional average of 
12.3. based on three specimens from Carico Lake 
Valley). In contrast, the three recognized iso- 
lated subspecies of Gila bicolor ( table 30, fig. 1 9 ) 
have a slightly higher average ( unweighted mean. 
16.40) than the populations retained in G. h. 
obesa (unweighted mean. 15.71 ). The respective 
ranges of means are 15.88-16.71 and 15.30- 
16.27. The means (13.48-14.17) are low in 
Reliclus (table 41). in comparison with those 
(15.30-16.71) in the Gila .samples. Thus, the 
trend toward reduction in meristic numbers in 
the i-solated spring populations, which are usually 
dwarfed, is only partially confirmed. 



88 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



-1 7.5 



O 
o 



70 







+ 


• 
• 
+ 

• 

+ 

• 




(!) 


-H- 




.i) (I) 
.1 







140 



125 130 135 

MEAN NO OF PECTORAL RAYS 

FiGLiRE IS. Positive correlation, between popula- 
tions ot Rhmuhiliys asciilus. m the mean niiinhers of 
pelvic and pectoral ra\s. Hntrics tor isolated endemic 
subspecies are circles; those tor populations trom the 
Diamond Valle\ system are crosses; those for popula- 
tions from the Humboldt River system (one introduced 
into Ruby Valley) are unringed black dots. Data from 
table 15. 

The pelvic fays niiniher 5-9 in tiie Rliinichtlns 
samples (table 15) and 6-1 1 in the Gihi pupula- 
lions (table 30), with a strong tendency toward 
reduction in modal number in each species in the 
races and subspecies confined to spring waters: 
from X to 7 in RliinichtliYs (fig. IS) and from 9 
to S in Gild (fig. 19). In Rrliciiis (table 41 ) the 
observed variation is 5-9, btit the mode is shaiply 
and consistently at 8. 

The mean numbers of pectoral and pelvic rays 
are strongly correlated positively between popula- 
tions in the pooled Rliiiiichlhys samples, pri- 
marily because there is a reduction in the means 
for each fin in the isolated spring subspecies (fig. 
IX). In the Gild populations, in direct contrast, 
there appears to be some interpopulation negative 
correlation between the pectoral-ray and the 
pelvic-ray numbers ( fig. 19). because the isolated 
differentiated races tend to have a slightly in- 
creased pectoral-ray count, along with a markedly 
reduced number of pelvic rays. No significant 
correlation is shown in this respect by Rcliclus. 
because there is little fluctuatiim in ray number 
in each fin. 



■ • 

\ 

•1 



150 155 16 165 170 

MEAN NO OF PECTORAL RAYS 

Figure ]'-). Low mean number of pelvic rays as- 
sociated with high means for pectoral rays in popula- 
tions of GiUi hicolor. Lone dots represent collections 
from Humboldt River system; crosses, those of Diamond 
Valle\ sxstem; encircled dots, isolated subspecies. Data 
from table }i). 



Within populations, however, the positive cor- 
relation between the numbers of pectoral and 
pelvic rays is at most extremely slight. For 
Rhiniclitliys a positive correlation (/• = 0.39± 
0.06) is very slightly suggested for the combined 
data for the populations of the Diamond Valley 
drainage (table (S). but the interpopulation cor- 
relation may be at least partly responsible. Simi- 
lar correlation tables for four subspecies of Gila 
hicolor { not here reproduced ) show a similar, 
extremely slight, far from certain, indication of 
positive correlation: ;— 0.26^0.13 for the 



Table S. ConcUilion hclwccu mimhers of i-ays in 
holh pelvic and hi'lli pcclcral jins in all four populations 
of Rhinichthys osculus m the Diiunond Valley ilroinage 
system (r = 0.39 ± 0.06). 



Total 
pelvic 
rays 






Total pectoral 


rays 






24 


25 


26 


27 


28 


29 


30 


31 


i: 


1 




1 




_ 




_ 




13 





— 


— 


— 


— 


— 


— 


— 


14 


10 


12 


?9 


9 


IS 


4 


1 


— 


15 


1 


— 


12 


5 


16 


1 


3 


1 


17 


1 


? 


?0 


12 


41 
1 


9 


13 
1 


I 


18 


— 


— 


— 


— 


1 


— 




— 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



89 



Table 9. BUateidl vaiialion in niiiulwr of rays in paired fins- of cyprinid fishes in llic Greai Basin in Nevada. 



Total 


Count hi 


igher 


Asymmetry 
index' 


Dextrality 
index' 




number 


Left 


Right 


100(L+R) 


100 R 




counted 


L 


R 


N 


L+R 


100 P' 


503 


51 


80 


26 


61 


I.I 


365 


55 


50 


29 


48 


62 


308 


50 


48 


32 


49 


84 


115 


15 


14 


25 


48 


65 


in 


47 


42 


12 


47 


60' 


929 


76 


51 


14 


40 


2.6 


409 


43 


15 


14 


26 


0.02 


1884 


182 


112 


16 


38 


0.00 


668 


73 


53 


19 


42 


7.5 


229 


20 


12 


14 


38 


16 



Pectoral counts 

Rhiniclilliys osculiis^ 
Gila hicolor 
Gila bicolor' 
Reliclus solitariiis-' 

Pelvic counts 

Rhinichtliys oscalns'' 
Rliiniclitliys oscahis' 
Gila bicolor pccliiiifer" 
Gila bicolor' 
Gila bicolor' 
Reliclus solitarius' 

1 Metliods of computation and presentation based on Hubbs and Hubbs (1945). 

-Original data, based wtiolly on material herein reported. 

"Data from Hubbs and Hubbs (1945. pp. 274-276); material slightly overlapping that listed in the next Ime. 

* Based wholly on material herein reported, partly overlapping that reported by Hubbs and Hubbs. 
^ Based on representatives from all basins. 

"Data from Hubbs ,and Hubbs (1945, pp. 276-277), based on a single collection from Walker Lake, Nevada; not included in material used for 
next two lines. 

* Value corrected for 100 P. 



Lovelock sample of G. b. obcsa, 0.27 ± 0.14 for 
the .sample of G. b. newarkensis from near Dia- 
mond Peak, 0.24 ±0.17 for G. b. citchiki, and 
0.27 ±0.19 for G, b. isolata. A low degree of 
positive correlation ( /• = 0.23 ± 0.09) is shown 
by the pooled data ( also not here reproduced ) for 
Relictiis solitarius. 

All of the data for the pectoral and pelvic rays 
in the material treated in the present report indi- 
cate very high positive correlation between the 
number of rays on the left and right side, with, 
as usual, a strong tendency for deviation from 
partner to increase in frequency with deviation 
from the mean value of the population, and to 
diverge from the partner in the direction of the 
mean value — all in line with the general rules. 

An analysis of asymmetry in the ray number 
of the paired fins (table 9), in the same material, 
indicates that among specimens with an asym- 
metrical count the average may be statistically 
higher on one side than on the other, in line with 
the exposition by Hubbs and Hubbs ( 1945: 
methods of calculation and expression on pp. 
232-233; comparable material on pp, 263-277). 
For the pectoral-ray counts, a slight tendency 



toward dextrality is indicated for the pooled data 
on all of the Rliiniclitliys osciiliis subspecies 
treated, but no significant deviation from sym- 
metry is evident for the subspecies of Gila bicolor 
or for Reliclus .solitarius. For all three genera, 
the data for pelvic rays indicate sinistrality ( that 
is, higher counts statistically the more frequent on 
the left side). The evidence is conclusive for the 
pooled data on Gila bicolor. but somewhat less 
so for Rliiniclitliys osculiis and not of high statisti- 
cal significance for Relictiis. Previous data, based 
in part on the same material ( Hubbs and Hubbs, 
1945, pp. 276-277), is confirmatory for Gila 
bicolor, but shows statistically untrustworthy 
sinistrality for Rliiniclitliys osculus. Possibly, the 
trend toward sinistrality may be somewhat race- 
dependent. The index of amount of asymmetry 
seems to be remarkably consistent for the three 
species treated, ranging in all three, for pooled 
data, from 12 to 18 percent for the pelvic-ray 
data and from 25 to 32 percent for the pectoral- 
ray data. The higher index of asymmetry for the 
pectoral fin probably reflects in part the higher 
absolute number of rays. 

The observations on bilateral asymmetry in 



90 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



ihc paired fins o\ all llircc species studied eim- 
toriiis uilli liie principle, fiist uxuked out and 
denuMislrated tor Lcptoccltiis armutiis ( Hiibbs 
and Hubbs. 1945. pp. 266-272. fig. 1). and 
since found lo hold uni\ersall\. for both pectoral 
and pelvic fins, that, when the ccnnits differ on the 
two sides, the lowest (innermost I and presumably 
last-formed ray of the fin with the higher ray 
number is shorter (and generalK weaker! than 
the corresponding ray on the opposite fin. C om- 
parison showed that with very few exceptions 
this condition holds for the fishes under treat- 
ment. For the counts that carried the pertinent 
notation on the data sheets, the lowest ray in the 
paired fin having the higher count was shorter 
than the lowest ray in the opposite fin. of seem- 
ingly equal length, or longer, in the following 
ratios: for the pectoral fin. S.V|:2 for Rliinicli- 
Ihvs. !?> ■.()■.] for Gila, and 16:0:0 for Rclicliis: 
and for the pelvic fin. 93:0:1 for Rhinicluhys. 
72:0:0 for iiihi. and 16:0:0 for Rcllciiis. The 
principle now seems to be soundly established. 
not only for numbers of rays in the paired fins, 
but also for numbers of parts in xarious other 
bilaterally paired nieristic structures. 

Vertebra!'. The general availability of high- 
grade X-ray equipment in ichthyological labora- 
tories is leading to the extensive use of \ei'tebral 
nimibers (and other ostetilogical characters) in 
fish taxonomy (Miller. 1957). To test for pos- 
sible differences in vertebral numbers, samples 
from each of the populations of each of the three 
species under treatment were X-rayed. C ounts 
were obtained and recorded for the total mmibers 
(tables 16. .1 1 . 42), including the entire hy|nual 
plate as 1 and the W'eberian apparatus as 4 (the 
filth vertebra, following that apparatus, was 
readily identified by its essentially normal centrum 
and strong neural spine). Because of variations 
in the length, strength, aiui arrangement of the 
hemal spines and the interhemals near the junc- 
tion, the distinction of precaudal and caudal 
vertebrae was perhaps somewhat subjective, and 
was abantlonei.1 for (Jilci ami Rclicliis. which, 
furthermore, shovsed no sharp differences in the 



ratio between the numbers in the two sections, but 
was retained for Rliiiiiclitliys. \\iv wiiich the dis- 
tinction seemed less difficult and some average 
differences appear for each section (table 16). 
Counts were disregarded whene\er the radiograph 
showed any marked abnormalities, or when (infre- 
quently in some populations), any of the neural or 
hemal spines were doubled on one or more of the 
piisteriormost caudal \ertebrae ( it might have 
been better, and in line with sc^me practice, to 
have enimierated the centra, without regard to 
any doubling of the spines ) . Within each species, 
niter-population differences in \ertebral number 
are rather limited: there is some overlap between 
all |iossible pair combinations. There is also a 
wide overlap in the total numbers hir the samples 
of each species: .v5 — 10 for Rliiiiirhlliys. 37 — 12 
for (Jihi. and 35-39 for Rcliciiis. The means are 
consistently lower than 3S.0 for Rliinichthys (ex- 
cept for the 3 specimens from C'aricti Lake Val- 
ley) and for Rclicliiy. but are higher than 3.S.0 
for Gild. 

Because coirclations. on both phenotypic and 
genotypic basis, have often been loimd between 
vertebral numbers and the en\ ironmental tem- 
peratme. and since the habitats of Rliinichthys 
<>\ciilii\ throughout the area studied vary from 
cold springs to hot spiings. the mean total nmii- 
ber of vertebrae has been graphically compared 
with the temperature of the water at the time of 
collecting (fig. 16). The temperature readings 
tluHigh isolated aie presumably repiesentative of 
basic differences, because all (except those for 
the samples from Dianas Punch Bowl ) were taken 
(.lining the warm |iart of the \ear in spring-fed 
waters. No e\ii.lent coirclations appear in the 
graph. Possibly, in reponse to adaptational ad- 
justments to .secure normal development at differ- 
ent temperatures, similar meristics have been at- 
tained in waters of c|uile i.lifferenl temperatures. 

For Gila, the mean vertebral numbers for the 
7 stations with recorded temperatures of 20.0 to 
23.3 C . ranged from 3S.5() to 39.86, with the 
two extremes at 20.0 C: the springs designated 
as "cold." ""cool." ain.1 ■"just under luke-warm" 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



yielded fish with averages of 39.07. 3S.39. and 
38.78. respectively; the Location with the highest 
recorded temperature did have the lowest verte- 
bral mean, 38.23, but this collection (Warm 
Springs in Independence Valley) harbored G. b. 
isohita. the most dwarfed of all the populations, 
and low counts are often enccnmtered in dwarf 
forms. 

For Relictus the mean vertebral numbers 
ranged from 36.05 to 37.08, with a slight indica- 
tion of lower values at the warmer Collections; 
5 means for Collections at about 12 to I 8 C. 
averaged 36.71; 2 means for Collections with 
temperature records ranging so as to overlap the 
two other sets averaged 36.46; and 3 means for 
Collections at 2 1 to 25 averaged 36.32. The 
slight correlation may be fictitious, for the two 
extreme temperatures were for Collections ( 7 and 
1 1 ) rather close together in the same drainage 
region. 

A similar plotting of average numbers of dor- 
sal, anal, pectoral, and pelvic rays of Rhinichlliys 
oscuhis (of all three subspecies) against the re- 
corded spring-water temperatures ( fig. 1 7 ) 
showed the same essential lack of correlation, 
nor was any correlation noted for the data on 
GiUi bicolor or Relictus soUhirius. it therefore 
seems probable that the local variations in ray 
numbers are also not attributable to the direct 
effect of the environmental temperature. 

Scale rows. A total of 12 different sets of 
scale counts were utilizxxl, 4 for Rliiniclilliy.s. I 1 
for Gila, and 5 for Rcliciiis. Irregularities of 
rows and the frequent incomplete development of 
the lateral line led to the deleting, for Rliinichthy.s 
and Relictus. of 3 sets of counts, for rows above 
and below the lateral line, and the total number. 
To replace the usual counts "above lateral line" 
and "below lateral line," counts were taken in 
Relictus between the origins of the dorsal and 
anal fins. In Gila only, scales were enumerated 
in a series, "lateral line to pelvic," running upward 
and forward from the pelvic-fin insertion to but 
not including the lateral line. 

The methods of counting are those specified by 



Hubbs and Lagler ( 1964, pp. 22-23 ). The "cir- 
cumference scale count" and "caudal pedimcle 
scale count" of that treatise are here abbreviated 
to scale rows "around body" and "around pe- 
duncle," respectively. In addition to the total 
count in each of these two sets, separate enumera- 
tions were made in Gila of the scales in those 
circumferential counts that lie respectively above 
and below the lateral line. The two lateral-line 
rows are added to yield the total-circumference 
count. 

In Gila, the scale rows are relatively regular, so 
that the enumerations are subject to little doubt 
or subjective interpretation. In some forms of 
Rhinichthys and in Relictus. however, some of the 
counts are approximations, because the scales are 
more or less deeply embedded, p<.)orly imbricated, 
and covered by mucus, and are small, often ir- 
regularly seriated, and subject to considerable 
loss and regeneration. Attempts to scrape off 
excess mucus often results in descaling the body. 
Undue fluctuations in counts by reason of such 
circumstances weie largely overcome by adopting 
appropriate magnification and illumination and 
by using a fine jet of compressed air. 

Most of the scale counts enumerated for Gila 
were niaele for us. using our methods, by a re- 
search assistant, John T. CJreenbank, in 1942-43. 
The more difficult counts, those for the two other 
genera, were all made by one of us (R.R.M.), 
to help assure uniformity. 

All three species vary considerably in scale 
counts. For the four sets of counts available for 
all 3 species, using pooled data (table 10). 
Rhiniciuhys is generally the most variable and 
Relictus seems to be the most constant (but fewer 
specimens of the latter were counted). 

There are some marked differences between 
populations of Rhinichthys osculus and Gila 
hicolor in numbers of scales in the several rows 
(tables 18 and 32). For example, the Indepen- 
dence Valley subspecies. Rhinichthys dsciilus 
k'lhoponis and Gila bicolor isohita. diffei' almost 
consistently from the other subspecies of each 
species in nearly all scale counts. Interpopulation 



92 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Tarlk Id. Ori'}-iill viiiiatioii in scale taunls of the S specifs of Cypiiniihw in ihc north-central (/real Basin. 

Range of values for populations counted 



Lateral-line series 

Rhinielithys oseiilns ^ 239 )' 
Gila hieolor ( 260 I 
Relictiis solilariiis 1 75 I 

F'rcdoisul rows 

Rliiiiichlliys osenlii\ ( 235 I 
Gila hieolor ( 249 ) 
Relieliis solilariiis ( 65 ) 

Around body 

Rhinielithys oseiilns (211) 
Gila hieolor (249) 
Relieliis solilariiis ( 65 ) 

Around pcdimclc 

Rhiniehlhvs oseuliis (215) 
Gila hieolor (249) 
Relieliis solilariiis ( 65 ) 



Minimum 



50-66 (17)" 
42-52 (II) 
50-59 (10) 

26-tO (15) 
22-28 (7) 
27-32 (6) 

4S-6() (13) 
40^51 (12) 
52-60 (9) 

26-32 (7) 
20-29 (10) 
28-32 (5) 



Mean 



56-71 (16) 

47-57 (II) 

54-63 (10) 

30^43 (14) 

27-31 (5) 

31-34 (4) 

54-67 (14) 

44-56 (13) 

55-63 (9) 



30- 
25- 
29- 



36 (7) 

32 (8) 

33 (5) 



Maximum 



60-75 
50-64 
55-70 



( 16) 
(15) 
(16) 



35-47 (13) 
29-34 (6) 
33-39 (7) 

60-76 (17) 
49-63 (15) 
58-66 (9) 

31-40 (10) 
28-35 (8) 
30-34 (5) 



^ These numbers in parentheses after the species names indicate tlie number of specimens counted for the stated scale row. 

- Numbers in parenllieses indicate tile range in counts for the mmimiim. mean, and m:i-\inuim values recorded for eacll species 



differences in Relieliis soliuiriiis in scale counts 
are less striising (p. 223. table 43). 

Lateral-line pore counts are discussed above. 

Gill-raki:rs. The raker count, often made 
on both sides represents the total number on the 
outer face of the first arch, including the few at 
and just above the rounded angle. With due 
care, the numbers can be ascertained with high 
precision and repeatability. We use appropriate 
magnification, adequate illumination, and a fine 
jet of compressed air. It is imperative that all 
rudiments at each end of the series be inchided. 
because slight rudiments grade into well developed 
rakers. In the cyprinids studied, unlike clupeids 
and some other fishes, we have found that speci- 
mens as short as 30 mm. had attained the full 
numerical complement of rakers, so that such 
small specimens could be safely counted as repre- 
sentative of the species. 

In the populations studied, the rakers varied 
widely in number. In Rhinielithys and Relieliis 
the wide variation, from 5 to 10 and from 7 to 12. 
respectively (tables IX, 43), is consistent with 
the principle that degenerate structures tend to be 
highly variable. The vastly greater total fluctua- 
tion, from K to 40 for the whole complex that we 
treat as a single species, GiUi hieolor. and the 



concomitant variation in size, texture, and struc- 
ture, is attributable to trophic divergence (pp. 
146-147). 

Counts only were considered for the popula- 
tions of Rhinieluhys and Relieliis, for in these 
genera the rakers are consistently few, short, 
fleshy, and weak, but for the ]ioiiulatioiis of 6'/7(; 
hieolor consideration was also given to degree of 
development. In that species, in apparent trophic 
adaptation, as in many species or species groups 
of fishes, the rakers diverge widely in size, texture, 
and structure (fig. 28), as well as in number. 
Some of these differences are appreciable by in- 
spection, and are subject to description. For 
quantification of length, we utilized the length 
of the longest raker ( usually readily selectable by 
inspection ), which lies slightly below, or rarely at, 
the angle. The left arch was consistently counted. 
The measuremeni has been obtained with preci- 
sion dial calipers marked in 0.1 mm. intervals 
and estimated to 0.01 mm., with an error of prob- 
ably less than 0.05 mm. One point of the calipers 
was pressed lightly against the, often more or less 
concealed, hardened base of the raker. Since a 
graph showing raker length plotted against 
standard length demonstrated an essentially 
straight-line relation between those parameters. 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



93 



the raker lengths were talHed as permillages of the 
standard length. Because of differences between 
races in proportionate head length, raker lengths 
were also tallied against head length. Between, 
and more loosely within, populations, there is, in 
Gila hicolor (.see p. 146 for details), roughly, a 
positive correlation between number of rakers and 
their size ( fig. 29 ) and between number and 
form. 

Another special aspect of the meristic analyses 
has been the determination of the correlation 
within the GiUi hicolor complex (whether we call 
these forms subspecies or species ) between num- 
bers of gill-rakers and of pharyngeal teeth ( p. 
145; table 24), as an approach to the analysis of 
introgression (pp. 144-147) between the lacus- 
trine form { pectiiiifcr) and the fluviatile form 
( ohcsa ). Much variation has been found in num- 
bers of both gill-rakers and teeth in populations 
of Gila bicolor that show evidence of past hybridi- 
zation between those forms (p. 145). 

Pharyngeal teeth. Numbers of pharyngeal 
teeth have traditionally been emphasized in the 
classification and nomenclature of cyprinid fishes, 
but few studies have assessed the degree of in- 
dividual variation or have critically tested the 
value of the observed patterns. In this study, 
more than the usual casual attention has been ac- 
corded to variation in the number and arrange- 
ment of these teeth. Large series of discrete pop- 
ulations have provided appropriate material for 
this study. 

In determining the number of the teeth, it must 
be kept in mind that individual teeth are fre- 
quently lost in the normal process of tooth replace- 
ment and sometimes in the extraction of the 
arches ( which calls for care, experience, and 
skill). Loss of teeth must be evaluated by rec- 
ognition of the alveoli, which need be differ- 
entiated from nerve and blood-vessel perforations 
(see fig. 45). Ordinarily, alveoli are larger, are 
rimmed, and follow the pattern of tooth arrange- 
ment. 

Difficulties in counting increa.se with the age of 
the fish: because in old fish the teeth are not 



always replaced, margins of the arch may become 
eroded, and the alveoli may become more or less 
closed with bony tissue. Younger fish seldom 
show such modifications, and are therefore pref- 
erable for counting, and for the study of arch 
and tooth structure. With good tools, care, and 
practice, arches can be extracted from even ex- 
tremely small fish, without damage to arch or 
body. 

One of the surprising and unexpected finds in 
the present study is that the number of teeth in 
the main (morphologically outer) row is occa- 
sionally reduced to ? in all three genera studied 
(pp. 97, 148, 190). It was long assumed that 
the only modern North American cyprinid having 
only 3 teeth ( in this case normally on both sides) 
is a peculiar genus and species, Stypocloti sii^nifcr 
Carman, which occupied an endorheic basin in 
northern Mexico but is now probably extinct 
(Miller, 1961, p. 380). The occasional reduc- 
tion of the tooth number to 3 on the right side 
of any North American cyprinid was apparently 
first recorded by Miller ( 1945a) in describing 
dental variation in Snydcrichthys (now Gila) 
copci (Jordan and Cilbert), in which a syntype 
of Sqiialiiis aliciae yielded a formula of 2, 4 — 3, 
2 and a specimen of G. copei from Little Wood 
River. Idaho, had a formula of 1, 4 — 3. 1. This 
deviation was also noted by Hubbs and Hubbs 
( 1958, pp. 299, 303). in the original description 
of another cyprinid of northern Mexico, Notropis 
saladonis. Overlooking the reported variants of 
G. copei, and also Stypodoiu Lachner and Jenkins 
( 1967, p. 577) stated that reduction to 3 teeth 
had been found, among North American cyp- 
rinids, only in Nolropi.s saladonis and in Nocomis 
leptoccphaliis (Girard); in the latter they noted 
occasional reduction to 3 on one or both sides. 
We have found the teeth in Agosia chrysogastcr 
Cirard to vary in one specimen from the normal 
complement of 4 — 4 to 3 — 4 ( an exception to the 
general rule of having the higher number of teeth 
on the left side; similarly, we have found in Gila 
bicolor, rarely, variants with 3 — 4 or 4 — 5 teeth ) . 
In Aztccula vittala (Cirard). from the Valley of 



94 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Mexico, rare variants from tiie 4 1 norm are 

3—4, 4—3. and 5—4 ( 1 each). 

La Rivers ( 1962, p. 427. fig. 196) figured a 
pharyngeal arch of Rliinichthys osciiliis itcvadcn- 
sis Gilbert as toothless and stated that the denti- 
tion of this form "varies from no teeth (() — 
— 0) to the following combinations — — 1 / 
3—0. 0—3 1 — and 0— 1 /O— 0." obviously on 
the basis of damaged or diseased arches. 

The occasional increase in the number of teeth 
in the main row beyond 5. to 6. was noted twice 
in a sample of GiUi hicolor obcsn. Evans and 
Deubler (195.S. p. 32) reported h teeth in the 
main row in a specimen of Semoliliis iitromacii- 
hiius (Mitchill). It has generally been thought 
that the number is increased beyond 5 in only one 
North American cyprinid genus. Orihodoii (of 
the Sacramento River system). 

Snyder (1917. p. fi2 ) noted that -'Sii>luilclcs 
ohesiis" (as lie called the stream form) usually 
has 5 teeth on the left arch and 4 on the right, 
and Hubbs and Hubbs (1945. pp. 2S4-2H5) 
generali/ed that those cyrinids that are bilaterally 
asymmetrical in pharyngeal-tooth number have 
the higher number on the left arch. Both prior 
and sub.sequent observations have solidified this 
conclusion. For other species of Gihi. Hubbs and 
Miller (1943. p. 356). Miller (1945b. p. 107; 
1963. p. 23). and Miller and Hubbs ( 1962, pp. 
I 12-1 13) have found the teeth in the main rovs 
to be almost always 5 — 4 (following convention 
in listing the left side first). Evans and Deubler 
(1955. p. 32) stated that all 150 specimens 
counted of Clinostoinus clongatiis Kirtland have 
2, 5—4, 2 teeth. 

Exceptions to the rule that the count on the 
left side, if different, is the higher, are provided b\ 
individual variants only. The only contrary state- 
ment found in the literature is that by Chu ( 1935, 
p. 107), who stated that "Pho.xiniis .... has the 
teeth on the right side constantly 2. 5 while those 
on the left side are 1 or 2. 4 or 5 (0, 1 or 2, 4 or 
5/5. 2, ()).'■ However, in the same treatise (p. 
116) he gave the formula as "2, 5/4 or 5, 1 or 2" 
for Pli<>\inu\ Idi^ow.skii V(irii'i;(itu.\ Gunther and 



figured (pi. 22, fig. 105) a right arch with 4. 2 
teeth. Plioxiiuis phoxhius (Linnaeus) is known 
to agree with other cyprinids in normally having 
the higher count on the left side ( Hubbs and 
Hubbs, 1945. p. 2X5 ). Specimens of P. phoxinus 
and P. pcrcnunis ( Pallas ) examined by us have 
5 — 4 teeth in the main row. Obviously. Chu's 
statement was a lapsus caltimi. 

In the material under treatment of all three 
genera and species, the higher number of teeth 
in the main row is on the left side. In Rhinichthys 
osciiliis the only variant formulae from the normal 
4 — i anuMig 79 specimens are 5 — 1 in two and 
4 — 3 in one. In Gila hicolor. among the great 
majority having bilaterally asymmetric counts, the 
only variants from 5 — I are 6 — 1 in two. 4 — 3 
in t)ne, and, as a leversal of laterality, 4 — 5 in 
only three fish (table 23, p. 149). In Relictus 
\oliliiriiis. among 75 specimens counted, the only 
variants from 4 — I are 4 — 3 and 5 — 4, each in 
two fish. 

The remarkable constancy in the number of 
pharyngeal teeth on each arch seems to be re- 
latable to the individual specialization of the teeth, 
which is indicated by the regular sequence of re- 
placement (Evans and Deubler. 1955) and by 
the distinctive size and form of the particular 
teeth. This distinctiveness was observed in all 
three species, but may be illustrated by the n^na- 
tions made for Gild hicolor. The first (upper- 
most and posteriormost ) tooth is long and very 
slender. In some specimens it is crowded to the 
outer side (that is. toward the concavity of the 
arch), where it is closely juxtaposed against the 
.second tooth. That tooth is usually also very 
slender. The third tooth is mLich more robust, 
more compressed, and often the longest. The 
fourth tooth is usually somewhat shorter, but, on 
the left side, is also robust and compressed. On 
the right side, the fourth ( and last ) tooth is usu- 
ally much shortened, but wider than the third, 
and the grinding surface on the fourth tooth is 
usually less developed than ou the preceding 
teeth, although this tooth usually retains the 
stronu hook. The fifth tooth, normallv formed 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



95 



only on the left arch, is almost invariably a small, 
pointed peg. well separated from the fourth tooth. 
Occasionally, however, it ends in a slight to 
moderate hook and, rarely, it develops a small 
grinding surface. 

For all three species represented in the study 
area, the data on tooth number is consonant with 
the generalization that the number of serial ele- 
ments becomes reduced in populations inhabiting 
isolated waters. Rliinicluhys osciiliis robiistus, 
which is largely confined to springs, normally has 
only one tooth in the lesser row (p. 97 ), whereas 
subspecies inhabiting integrated streams often 
have two teeth in this row. Gilii bicolor ohesa and 
its derivatives, which usually occupy springs and 
.small creeks, normally have a dental formula of 
5 — 4, whereas the lacustrine type, G. h. pectiuijer 
(Snyder), typically has 5 — 5 teeth. Relictiis 
solitarius. which is virtually confined to springs, 
normally has 4 — 4 teeth. 

The form of the pharyngeal arch has also been 
accorded attention, particularly in the generic 
comparisons (pp. 189-193). 

Distribution and Habitat. 

A major phase of the study of the fishes of the 
isolated waters of the arid American West, 
specifically of the north-central Great Basin, has 
been an analysis of the distribution of the remnant 
fish populations in terms of paleohydrographic 
history (pp. 70-79). The problems of faunal 
depauperation, limited sympatry. and correlation 
between size of area and richness of the fauna arc 
summarized above. Particular distributions of the 
four genera under treatment, and of the repre- 
.sented species, subspecies, and races, are treated 
below under the respective headings. 

Treated below also, as a characteristic attribute 
of the several forms and populations, are their 
respective types of habitat. For each location, 
attention is paid to the isolation and the size of 
the habitat; to the abundance of the individual 
populations, largely in terms of numbers of speci- 
mens collected; to the temperature of the spring 
waters, which tends to be a critical factor ( most 



readings were in the Fahrenheit scale, which have 
been computed to the nearest degree Celsius); to 
various other major ecological criteria, such as 
clarity and other qualities of the water, and the 
nature of the bottom, current, and vegetation; 
and to associated fish .species, native and intro- 
duced. Listed also, when available, is the type 
of collecting gear ("woven-mesh seine" usually 
refers to the '"Common Sense" type ) . 

Unfortunately, little of the habitat information 
was quantified by us, and very little was recorded 
on the widely varying content of dissolved .salts, 
which presumably has been a factor of very con- 
siderable importance in the survival or extinction, 
and probably in the speciation of the populations; 
hence in their occurrence and distribution. It is 
unfortunate that systematic analyses have not 
been run, and studied in relation to the dwindling 
fish fauna. Prior to undertaking the major expedi- 
tion of 1938 the senior author made arrangements 
to have such analyses run in the pollution and 
water-quality laboratory operated at the Univer- 
sity of Missouri by the U. S. Bureau of Fisheries. 
Water samples were taken at the many springs 
examined and were dispatched at intervals to that 
laboratory, but no report was received. Fortu- 
nately, a considerable number of analyses have 
been published for spring waters in the basins 
under study, for example by Clark and Riddell 
( 1920) for springs in Steptoe Valley, and in the 
reports of the Nevada Department of Conserva- 
tion and Natural Resources providing water- 
resource appraisals of springs in many of the 
valleys of the state ( these reports are referred to 
in the preceding treatment of the remnant waters 
of individual basins and/or in the following ac- 
counts of the habitats of the several species). It 
is to be hoped that in future explorations of the 
area the chemical constitution of the isolated 
waters, and their other characteristics, will be still 
further analyzed. 

Outstanding features of the habitats of native 
fishes in the Great Basin are the vast reduction 
of surface water in which any fish could survive 
and the limitation of the fish to only parts of the 



96 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



iiiiiiisciile supply. In general, the native fishes 
are confined to valley springs, most of which issue 
along faults margining the valleys. Though 
many of such springs are extremely small, at times 
with virtually no discharge, the water supply tends 
to be more or less constant, and the very few 
fishes that have survived, in a small percentage of 
the basins, have become adapted to withstand the 
precarious habitats. 

The valley springs have been favorable for the 
survival of remnant fish life not only t)n ecological 
grounds but also in terms of k>ng perpetuity. Long 
existence is indicated not only by the very circum- 
stance of present occurrence, but also by other 
evidence, such as the extensive accumulation 
about siMiie springs of travertine deposits and of 
Indian artifacts, often of varied type and different 
degree of patination. 

As is repeatedly mentioned in the preceding 
discussions of the remnant fish life for each basin, 
and in the subsequent accounts of the species, the 
rather numerous mountain springs and creeks in 
some of the basins are devoid of native fish life. 
Introduced fish, especially trcnit, may multiply 
and persist, at least for some years, until the 
hazards of existence become insuperable. In 
years of great drought the mountain-side and 
canyon-bottom springs and the canyon streams 
may become temporarily dry. FLuthermore. they 
are subject to the occasional, even though gen- 
erally very infrequent, fate of violent scouring by 
the torrential precipitation that characteriz.es the 
desert climate. The fishes are washed out onto 
the playas. and the waters then subside too 
abruptly and too completely to allow the fish to 
return to habitable waters. 

Evidence of diminishing waters — in lakes, 
streams, and springs — strike the eye throughout 
the Great Basin. The postpluvial changes in 
climate have obviously been a dominant factor, 
but indications seem to be increasingly emerging 
that spring flow has been diminishing even with- 
out evidence of climatic change. The vast amount 
of water that accumulated in aquifers when the 
pluvial lakes filled large parts of the basins must 



have continued draining out by springs even after 
the present climatic conditions became estab- 
lished, and probably even during times that have 
been drier than the present. The wonder is that 
even a few fishes have survived in the precarious 
habitats of the Great Basin. 

Struci UKi-: AND Stai us OF Populations. 

Attention has been paid to the structure and 
status of the various localized populations of the 
three cyprinid species that are native to the north- 
central Great Basin. Particularly for Rhinichtliys, 
the mass random collections have been analyzed 
to show the distribution of the fish by size and 
length. It is concluded that Rhinichtliys. and 
probably the two other genera, ordinarily spawn 
in the summer, and that for some weeks the young 
of the two sexes are about equal in number and 
in size. As yearlings, females are more numerous 
and somewhat larger than males, and comprise a 
much larger biomass. After spawning as yearlings, 
the males seem to die off or to stop growing, 
whereas some lemalcs appear to live on and con- 
tinue to grow . 

The past, present, and possible future of the 
populations have also been given consideration. 
Extensive local questioning, fortunately during the 
time when observant early settlers were still alive, 
has yielded evidence on the indigeneity or intro- 
duction of species, and on changes in population. 
The evidence is recounted under the headings of 
the several foinis. 

SPECKLED DACE 
Rhinichthvs osculus (Girard). 

This species, one of the four that have persisted 
in the complex of basins under treatment, is the 
most ubiquitous freshwater fish in western United 
States. Uncounted local variants occupy an al- 
most endless number of habitats, ranging from 
torrential creeks in the major river systems to the 
tiniest and most isolated spring holes in the 
desiccated valleys that comprise the Great Basin. 
Hints of this situation, particularly as it applies 
to the remnant waters of the arid West, have been 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



97 



given by us ( Hubbs and Miller. 1948b), and 
others, but the very magnitude of the problem 
has retarded extensive critical studies. Many of 
the forms are highly localized. For example. 
RhiiiicJithys osciilu.s tlwrmalis (Hubbs and Kuhne, 
1937 ) is known from a single warm spring in the 
Green River basin in Wyoming. 

Varying generic treatment has been accorded 
this species: during the present century it has 
been referred successively to Agosict, to the more 
restricted taxon Apocope, and, currently, to 
Rliinicluhys. That genus had previously been 
restricted to R. cilraliiliis (Hermann), which is 
widespread through eastern United States, and 
to R. ciilcimcttie (Valenciennes), which ranges 
across northern North America from coast to 
coast, and its derivative, R. evcniuinni Snyder, of 
the Umpqua River system in Oregon. Rliiiiiclitliys 
seems to be the best resting place for this western 
group, in the present confusion on generic rec- 
ognition among American cyprinids that appears 
to approach potential chaos. For the present, we 
follow Gilbert (1893, p. 229) in referring the 
group to Apocope Cope on the subgeneric level. 
The supposed generic distinction of Apocope on 
the basis of the continuous rostral groove breaks 
down, particularly in the Colorado River basin, 
where in many forms the premaxilla is commonly 
or even regularly bound down by a frenum. 
Agosia. which has usually been thought to be a 
closely related genus, occupying Williams and 
Gila rivers of the lower Colorado River system 
and tributaries to the Gulf of California farther 
south, may not be on the same phyletic line and 
we feel that it should be held separate. Except 
for two forms of the Columbia River fauna. R. 
falcatus ( Eigenmann and Eigenmann) certainly, 
and R. iimatilla (Gilbert and Evermann) prob- 
ably, we currently treat all the many named and 
still more numerous yet unnamed forms as con- 
stituting a single highly varying species, R. oscii- 
liis. Perhaps R. falcatus should be separated 
gcnerically, for it is trenchantly distinct. 

Despite the wide range of variation displayed 
in many characters, in large part related to the 



type of habitat, and in part to degeneration in 
isolated springs, the Apocope complex is held 
together by a considerable number of morphologi- 
cal and other features. The multitudinous local 
forms are all small minnows, rather slender and 
terete, with the body heaviest well forward, with 
rounded contours, and with a small eye and small 
mouth. They are generally rather dark and more 
or less striped and mottled, and, as the vernacu- 
lar name indicates, are speckled, because of the 
blackening of regenerated scales (p. 210). They 
have rather slimy integuments, and the deeply em- 
bedded, more or less irregularly aligned scales 
are smoothly oval, with no sharp distinction be- 
tween anterior, lateral (dorsal, ventral), and pos- 
terior fields, and with radii well developed around 
the entire scale (except in R. falcatus). There 
is no horny sheath on the jaws. The lower jaw 
is included, and the posterior tip of the mandible 
is not prominent. The small barbel is terminal or 
subtcrminal on the upper jaw, but is subject to a 
varying degree of obsolescence, especially in iso- 
lated spring habitats ( as is illustrated by the forms 
herein described as new subspecies ) . The pharyn- 
geal teeth normally number 1 or 2, 4 — 4. 1 or 2, 
but in the Great Basin area seem to be subject to 
little deviation from the formula 1, 4 — 4, 1. One 
variant from this formula, with 0. 5 — 4, was re- 
ported for R. o. robust us from Soldier Meadows, 
Nevada (Hubbs and Miller, 1948a, pp. 16, 17). 
Schultz and Schaefer ( 1936, p. 3 ) indicated that 
a considerable number of specimens of Rhiuich- 
thys osculiis lack any teeth in the lesser row. but 
we suspect that they overlooked some alveoli that 
represent lost teeth. Taking such alveoli into ac- 
count, we have found, for 79 specimens repre- 
senting all but one of the populations of Rhiuich- 
thys osculus herein treated, the formula is 1 . 4 — \. 
1, with these exceptions: 1. 5 — 4, 1 in two 
specimens; 1, 4 — 3, I in one; 2, 4 — 4, 2 in three, 
and 0, 4 — 1, in two. Other subspecies com- 
monly have two teeth in the lesser row. The 
ordinarily single tooth comprising the lesser row 
in Great Basin forms is almost always rather 
strong. The teeth are well hooked and the grind- 



98 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



iiig surtace is sometimes slightly cultrate and 
crenate. The aieh is relatively strong and sharply 
eiirved; its upper arm is weak and very flat. The 
gill-rakers eonsistently remain few and short, with 
no tendeney to increase m number or in length 
in correlation with feeding on plankton — such as 
has evolved in many cyprinids (and other fishes) 
throughout the world ( Hubhs. 1441b, pp. 187- 
ISS). The intestine is short, with a single simple 
loop in the form of a compressed S, equivalent to 
the Group 1 type of Kafuku ( 195S, fig. 7 and pi. 
4, la). The dorsal fin originates somewhat be- 
hind the vertical from the insertion of the pelvic. 
Typically, the dorsal rays number S (except in 
the Columbia River system ). the anal rays almost 
invariably 7, and the pelvic rays either 7 or 8. 
In males, the pectoral fin is enlarged in 
length and width and tends to be arched 
downward where the several outer rays are 
markedly thickened, so that the fin is reflected 
outward and downward along the outer (upper) 
edge (these characters are exaggerated in breed- 
ing males). The nuptial tubercles (fig. 2^H) 
tend to be obsolescent or obsolete on the head 
and body ( not so true in some forms of the Colo- 
rado River system), and on the fins are confined 
to the first several branched pectoral rays, on 
which the small hooks occur I per segment, in a 
single row, weak basally, but strong where once 
branched. They are diagnostically lacking on the 
much thickened first pectoral ray, on all other 
fins, and on the body ( not so restricted in some 
other subspecies). Though inhabitmg a wide 
range of habitats, particularly in respect to cur- 
rent, the varied forms are ordinarily mainly re- 
stricted to smaller waters (creeks and springs) 
that are neither very highly mineralized nor very 
warm. Furthermore, they generally avoid lakes, 
and any pelagic or deep habitat. None of the 
forms here under treatment have i.le\eloped en- 
larged falcate fins and the very slender caudal 
peduncle that are exhibited by certain local forms 
that have become adapted to life in consistently 
rajiid water (as is particularly well exemplified 
in certain tributaries of the Colorado River). 



Many of these characters seem rather superficial 
or even subjective, but together they put a stamp 
on the fish that makes it ordinarily possible to 
recognize it at a glance as belonging to Apocope, 
despite great variations in several respects. 

In so far as checked, the osteological charac- 
ters throughout the species essentially agree with 
those assigned to Rhinichthys in the account of 
the genus /^(7/(7//.s ( p|-). I S 1-1 9.^, figs. .■^8-4.'>). 

Included in the subgenus Apocope are various 
forms that until the I9.^0's were accorded specific 
rank. For example, Jordan and Evermann ( 1 896, 
pp. .'^()8-.i 1 ^ ) recognized in the subgenus 
Apocope the following species (referred ge- 
nerically to Agosia): Agosia osciihi. A. ydirowl. 
A. coiiesii. A. (ulohe. A. ncvciclensis, A. niihiUi. 
A. ctirringloiili. and .-f . vclijeni as well as A. 
iimntilhi and A. lalccitn. All of the forms other 
than Rhiniclil/nw iniuitillii and R. falcaliis we 
have long regarded as conspecific, and for 
the species we early adopted the name (^senilis. 
one of three proposed by Girard in 1 S.Sd (p. 186) 
as Argyreiis osctihi.y. A. not(ihili.\. and A. nuhilus. 
The name osciilus was definitely selected over 
notahilis by .lordan and Evermann (1896, p. 
.^M)9), by synonymizing Argyreus notahilis under 
Agosici o.H-iila. and was formally selected over 
iiidiiUi by Schultz (19."^6. p. 148). through the 
adoption of the names Apocope osciilci inihila 
and Apocope o.\cul(i ouiila. In temporary re- 
version, following the then accepted but now in- 
validated principle of the precedence of page and 
line reviser action over that of selection by first 
reviser. Miller ( 1952. p. M)) adopted Rhiniclitliys 
luiliiliis in jilace of R. o\ciiliis as the species name. 
The binomen Rhiniciuliys osciilus has been 
adopted by Bailey ct al. (1960. 1970) for all 
forms of Rhiniciuliys other than R. utralulus, R. 
CdlardcUic. R. cvcniuinni. an(.l R. jalcalus. 

Local Forms Of Rhimchthys oscui.us 
In NoRin-C 1 ntral Gri:at Basin 

In the basin area of Nevada under investiga- 
tion, local forms referred to Rhiniclithvs osciilus 
occupy five springs in the rather extensive pluvial 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



99 



Lake Diamond drainage basin, a single spring 
in the pluvial Lake Gilbert basin (recently ex- 
terminated ). and three springs in the pluvial Lake 
Clover basin. They also inhabit headwater creeks 
and springs in the surrounding drainage systems: 
to the north, the Columbia River; to the east. Lake 
Bonneville; to the south, the Colorado River; to 
the southwest, the isolated Lake Toiyabe basin 
(see p. i 5 ) ; to the west, the Reese River system. 
tributary to the Humboldt; and. to the northwest, 
the basins of the reputedly pluvial lakes Carico 
and Crescent, which in high flood discharge into 
Humboldt River. In the present study we defer 
detailed comparisons with the local forms that 
occur to the east, .south, and west, but have in- 
cluded four samples from the Humboldt River 
system immediately to the north ( including the 
two valleys just mentioned), as these seem to 
represent, as nearly as is feasible, the ancestral 
type that gave rise to the nine native populations 
discovered in the area of special study. These 
nine populations occur in three of the six basins 
that form, within the area of special study, the 
northern file of now completely enclosed depres- 
sions that were once connected with Humboldt 
River. Rhinichlhys osctilus is not represented in 
the basin of pluvial Lake Newark, nor in the 
basins of pluvial lakes Franklin (except by recent 
introduction). Gale, Waring, and Steptoe. all of 
which are occupied by only one native fish, the 
ecologically rather similar relict dace, Rclictus 
solitarhis. 

Altogether, huiuding extraneous samples for 
comparison and the one introduced stock, 14 pop- 
ulations referred to Rhinichtliys osculiis have 
been analyzed in the present study. They are 
here, as on the maps (figs. I. .^. <S. 12), referred 
to as representing Locations RO to R12. The 
series are referred to 4 subspecies, as follows: 

(A) Rliinichl/iy.'i osculiis robuslus (Rutter): 

From nearby basins discharging in rare floods into 
Humboldt River: 
RO, spring in Carico Latie Valley. 
Rl. Indian Creek in Crescent Valley. 
From present headwaters of Humboldt River, near 
Wells: 



R2, Bishop Creek. 

R.^. spring near Town Creek. 
Regarded as having been stocked from a Humboldt 
River headwater: 

R4. spring in Ruby Valley. 
From springs in basin of pluvial Lake Diamond 
(not differentiated enough for siibspecific 
separation ) : 

1^5. hot spring at Potts Ranch, near pluvial 
Monitor River, below Lake Diana. 

R5A. hot-spring flow 6 km. south-southwest of 
R5. at approximate source of any flow of the 
south branch of Stoneberger Creek. 

R(i. spring flow in Coils Creek (on Three Bar 
Ranch), a northern tributary of floodwater 
course representing Monitor River. 

R7, spring at Birch Ranch, on east side of the 
bed of ancient Lake Diamond. 

R8. Big Shipley Spring on west side of bed of 
same lake. 

Extensive inquiry and exploration has disclosed 
no other populations of the species in the 
basin of Lake Diamond, though some may 
exist in the immediate vicinity of Big Shipley 
Spring and/ or along the fault to the south- 
ward: and there is a report of fish thought 
not to be young trout in Roberts Creek (p. 
19). 

(B) Rhinichlhys osculiis reliquus Hubbs and Miller, a 

very distinct relict (recently exterminated): 
From basin of pluvial Lake Gilbert: 

R9, a single spring complex on Grass Valley 
Ranch. 

(C) Rliiiiichlhys osculiis olii;oporus Hubbs and Miller: 
From Clover Valle\. in the basin of pluvial Lake 

Clover: 

RIO. a large spring at Wright (formerly Ralph) 
Ranch. 9.5 miles south of Wells, near north- 
west tip of the ancient lake. 

Rll. large spring at Warm Spring Ranch, in 
southwest corner of the old lake bed. 

(D) Rhinichlhys osculiis Icihopoiiis Htibbs and Miller: 
From Independence Valley, also in the bed of 

pluvial Lake Clover: 
R12. Warm .Springs, a cluster on the west side 
of northern arm of Independence Valle\. 

In the separation of the subspecies, particular 
stress is accorded the degeneration of the barbel 
and of the lateral line. The development of these 
sensory structures in the forms of Rhinichtliys 
osculiis under treatment is now discussed. 



1 00 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 11. Niinihcr and size of htirhels in populations- of Rhinichthys osculus //; certain l^asins in Nevada. 



Subspecies 

Pluvial lake system 
Locality 





No. of barbels 






0—1. 




0—0 


1-0 


1—1 


1 


1 


1 


13 


23 


52 


16 





21 


11 


10 


19 


2 


2 


8 


12 


11 


lU 


6 


1 


13 


13 


9 


18 


42 


10 


9 


42 


41 


46 



Size of barbel 
(subjectively appraised) 



Absent 



Minute 



Small 



Large 



Rliiiiiclilliyx osciihis roblisliis 
L. Lahonlan (Humboldt R. ) 
Carico L. Valley 
Crescent Valley 
Bishop Creek 
Springs near Town Cr. 
L. Franklin (introduced) 

Ridiy Valley 
\ . Diamond 
Potts Ranch 
Dianas Punch Bowl 
Coils Creek 
Birch Ranch 
Big Shipley Spr. 
Rhiiiiclillir.s mciilns icliqiiiis 
1,. Gilbert 
Grass Valley 
Rliinichllixs o.sciiliis oliiiopoiiis 
L. Clover 
9.5 mi. S. of Wells 
Warm Sprs., Clover V. 
Rliinicluliys osculus lellwiporus 
L. Clover 

Independence Valley 



199 



25 
26 



75 



3 


— 


3 


— 


49 


42 


72 


13 


34 


9 


26 


9 


32 


28 


17 


3 



35 


18 


13 


— 


13 


5 


">■") 


— 


35 


8 


28 


9 


94 


17 


II 


— 


125 


55 


77 


1 



399 



50 
52 



154 



Development Of The Barbel 

In the study of the populations ot Rliinicluliys 
o.sciiliis ttnder treatment, it was found tiiat the 
barbel often requires close examination to be 
detected, because it is often lacking, extremely 
minute, or obscured from external view behind 
the concealed tip of the maxilla. It usually stems 
from the lower, far-posterior, outer surface of the 
maxilla, but in .some individuals is on the pos- 
terior edge, or hangs from the lower free edge of 
the maxillary fold, where it may be deeply con- 
cealed. Reliable determination recjuires adequate 
magnification and illumination, often supple- 
mented by the use of a fine jet of compressed air. 

As Snyder ( 1917. p. 67) indicated, the develop- 
ment of the barbel is subject to much variation 
between and within populations. As we have 
pointed out. the barbel, although commonly re- 
garded as a generic character of Rhiiiicliihys. is 
subject to obsolescence in local populations that 
are restricted to isolated springs. That this is not 
the direct phenotypic result of living in springs 



is suggested by the retention of the usual number 
and size of barbels in a stock recently established 
in a spring in Ruby Valley as an obvious result 
of introduction from a headwater tributary of 
Humboldt River (p. 107). In tho.se headwaters, 
as well as in the five populations that have per- 
sisted in the Lake Diamond system, which we also 
refer to R. o. rohiislns, the barbel is very fre- 
quently retained on one or biith sides, though 
with considerable variation ( table II). The four 
other populations that have persisted in the basin 
complex under treatment, and that we refer to 
three distinct subspecies, always or nearly always 
lack the barbel on both sides. These subspecies 
are R. o. rcHqiiiis of Grass Valley, in the basin 
of pluvial Lake Gilbert, and R. o. oligoporus 
and R. o. Icthoponis. of Clover and Independence 
valleys, respectively, both in the basin of pluvial 
Lake Clover. The last two subspecies, but not 
R. o. rcUquus. are further characterized by a 
great reduction of lateral-line pores (see below). 
In R. o. rcliqniis only one minute barbel, on 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



101 



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102 



CALIFORNIA ACADEMY OF SCIENCES 



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VOL. Vll HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



103 



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\o o 



— a ■ 



i;5D£ 



104 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



one side, was detected among 10(1 specimens that 
were ciiecked on both sides. In neither race of 
R. o. oligoporu.s was any trace of a barbel found 
on either side, among 51 individuals. In R. o. 
lethoponis, among 75 specimens examined, we 
detected, on one side only, a minute barbel in 
three and a small barbel in one. 

Rutter ( 1903. p. 148) described the barbels of 
R. (>. rohiisius as "usually absent, present on \0 to 
50 percent of specimens from any o\^c locality." 
Snyder ( 1 9 1 7. p. 67 ) . who gave definite numbers, 
indicated a rather similar range for stream popu- 
lations throughout the Humboldt system.' They 
may well have been dealing in part with local 
forms worthy of subspecific status, and may well 
have overlooked some rudimentary barbels. 

Development Of The Lateral-line Pores 

(.Sec also p. SI; tabic 12; tigiircs 25, 2h. ) 

in most of the populations of Rliliilchiliys 
osciilu.s here treated, the lateral-line pores on the 
body begin to form at a standard length of about 
22 to 25 mm. They increase in number slowly 
at first, then increase rapidly at the length of 
about 25 mm., and begin to approach the final 
number at the length of about .^5 to 40 mm. 
Relatively few specimens develop a full comple- 
ment of pores, though completion is approached 
in a few populations. With age, the lateral line 
becomes well tiver half complete at the Locations 
listed above under R. o. lohusliis. The popula- 
tions do differ somewhat in the rate of pore de- 
velopment ( note, for example, the different rates 
illustrated on figure 26 for the dace from Potts 
Ranch and Coils Creek, from two Locations in 
the Monitor Valley branch of the Lake Diamond 
system). The fish from Crescent Valley develop 
an unusually large number of pores (fig. 25), 
probably because they attain an Luiusually large 
size and have many scales (table LS). But high 
scale number did not yield so many pores in the 
fish from Carico Lake Valley, and accompanied 
one of the lowest pore numbers in R. o. icliqmis. 



' See SupplcnicnUiry Note (p. 253). 



Lahontan Speckled Dace 

Rhinklithvs osculus robustus ( Rutter). 

(Figures 22 aiul 23.) 

Pending the much needed and long deferred 
general review of the multitudinous local forms 
of the subgenus Apocope (defined above), we 
tentatively designate by this name the speckled 
dace of the Lahontan system and of certain basins 
that are part of that drainage complex, or almost 
certainly once were. In so doing, we follow Sny- 
der (1917, pp. 67-69), who provisionally 
adopted the name Ai^osia rohusta for the Lahon- 
tan representatives of "Agosia" although he rec- 
ognized the high variability of speckled dace 
throughout the Lahontan system and questionably 
separated the Lahontan complex of forms from 
those in other major drainage systems. Snyder 
may have included separable subspecies (ordi- 
narily, he did not recogni/e local forms within a 
stream basin, because of an obvious preconcep- 
tion that Western freshwater fishes are segregated, 
as full species, by major stream systems). Our 
decision to adopt the name robustus is based in 
part on the close agreement of our material from 
Carico Lake and Crescent valleys and from the 
headwater Bishop and Town creeks, and else- 
where in the Humboldt River system, with Rut- 
ter's type figure and description ( 1903, p. I4S, 
1 fig.) of Agosia lobusta. from near the western 
rather than the eastern edge of the wide Lahontan 
system. Rutter's later reference ( 19()S, pp. 139- 
140) to Agosia rohusta of populations from vari- 
ous parts of the Sacramento River system is ques- 
tionable, for more than one subspecies may well 
have been involved. 

Points of agreement between our material from 
the Humboldt River system and Rutter's type de- 
scription and figure include the rather faint and 
somewhat disrupted darkening of the primary 
dark lateral band and the weakness of the ventro- 
lateral band; a definite though only moderately 
strong continuation onto the snout of the blackish 
lateral band; the rather extensive pigmentation 
of the lower sides; the usual lack or at most weak 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



1U5 



development of horizontal dusky streaks along 
scale rows on side of trunk; the only moderately 
conspicuous blackening of the basicaudal black 
wedge: the blackening of the crotch of the rays to 
form a row of dashes near the middle of dorsal 
fin, and often, as a trace, in another row farther 
out on the fin; the only moderate speckling of the 
body; the frequent extension, though with inter- 
ruptions, of the lateral line well toward the base 
of the caudal fin; the only moderate obliquity of 
the nearly straight mouth, with a rather thin and 
strongly blackened upper lip; the rather small 
size of the head; the similar, not depressed, form 
of the snout; the small si/e and rounded form of 
the fins; the insertion of the pelvic fin well before 
vertical from dorsal origin ( on the average mid- 
way between base of caudal and a point just in 
advance of tip of snout ) ; the rather robust form 
of the body anteriorly, becoming moderately 
attenuate behind; and the similarity in body pro- 
portions in general (table 13). 

The Lahontan speckled dace has been treated 
as Rliinichtliys osciilus robiistus by Hubbs and 
Miller (1948a, pp. 17. 19, 22-24, 28; 1948b. 
pp. 44. 102); Shapovalov and Dill (1950. p. 
386); Shapovalov, Dill, and Cordone ( 1959. p. 
172); and La Rivers ( 1962. pp. 21. 89. 430- 
432 ) . The name was given as Rhinichthys imbihis 
rohustiis by La Rivers (1952, p. 99) and La 
Rivers and Trelease ( 1952, p. 117). 

It is possible that an older name, based on ma- 
terial from some other stream system, may be 
found applicable to the common form of the 
Lahontan system. Certain of these extralimital 
forms seem separable ( some perhaps even on the 
species level ) . Specimens from the Northwestern 
coastal area, which apparently should be called 
R. o. nubilus (Girard), have a much stronger 
dark lateral band (Schultz, 1936, p. 148). Col- 
lections from the upper and middle parts of the 
Colorado River system, representing R. o. yarrowi 
Jordan and Evermann and a chain of more or 
less sharply distinguishable forms from various 
tributaries, commonly to usually have a pre- 
maxillary frenum and otherwise look unlike R. o. 



robiistus. In other drainage systems the frenum 
appears to be consistently lacking. In the Great 
Basin area under special treatment, for example, 
this was found to be true: most of the many 
specimens checked for presence or absence of the 
barbel, whether they came from the Humboldt 
River system or from the isolated basins, were 
simultaneously checked for presence or absence 
of a frenum, without finding any. 

Rhinichthys o. robiistus also seems to contrast 
with various populations that inhabit much of the 
area, largely in Utah, that during pluvial time 
drained into Lake Bonneville, immediately east 
of the area under detailed treatment. For ex- 
ample, a cur.sory examination shows that the 
Lahontan type, and its derivatives, differ sharply 
from the form inhabiting springs along West Deep 
Creek. 6 miles above Ibapah, in White Pine 
County, Nevada ( near the midwestern margin of 
the Bonneville drainage, immediately east of our 
study area). Specimens from there (UMMZ 
141410), probably representing R. o. adobe 
(Jordan and Evermann), differ from R. o. 
robiistus in having a larger head; a lower, more 
nearly horizontal, and much larger mouth, with 
fuller lips, the upper of which is almost wholly 
devoid of pigment instead of being blackened; a 
larger and flatter snout; the lower sides barely 
marked by the second dark band, but generally 
darkened by horizontal dusky streaks along the 
scale rows (very seldom seen in the R. o. robustus 
series); a larger and more deeply blackened basi- 
caudal wedge, and a much more definite black 
streak on the snout. 

Rhinichthys o. robustus contrasts strongly with 
the forms herein described as distinct subspecies 
from basins adjacent to the Humboldt River sys- 
tem, and slight differences exist between the fish 
from the Humboldt headwaters and those com- 
prising the remnant populations in the Lake Dia- 
mond system. 

It seems to us highly probable that the popula- 
tions of Rhinichthys inhabiting the isolated basins 
herein under special study were derived from 
R. o. robustus or an immediate ancestor. In this 



106 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



view wc hold some reservation in respect to the 
recently exterminated form of Grass Valley, which 
we somewhat hesitantly restrict to subspecies 
status, as R. o. rcliquiis: it may be a specifically 
distinct relict of ancient and somewhat uncertain 
origin. 

Populations of Humboldt Rivlr System. 

(Figures 1, 12.) 

Four populations in Nevada have been selected 
to represent as nearly as feasible the race(s) of 
Rliinichthys osciihis that may be regarded as 
having originally occupied the basins of pluvial 
lakes Gilbert, Diamond, and Clover, wherein 
populations of this species are now rigidly re- 
stricted lo very isolated springs, and wherein, 
especially in the Gilbert and Clover basins, the 
populations have become markedly differentiated. 
Two of these four populations are those of a spring 
in Carico Lake Valley and of Indian C reek in 
Crescent Valley, both of which are in lake valleys 
that during high flood discharge into Humboldt 
River. The other two populations arc those of 
Bishop Creek and of springs adjacent to Town 
Creek, both near the town of Wells in eastern 
Nevada. These two latter Locations are in the 
perennial headwaters of Humboldt River, close 
to the divide that separates the Lahontan and 
Bonneville watersheds. These four Locations are 
designated RO for the Carico Lake Valley spring 
and RI for Indian Creek ( fig. I ) and R2 and R3. 
respectively, for Bishop and Town creeks (fig. 1 2). 

Loc:ilion KO. — Unnamed warm spring in (Carico 
Lake Vallcv. in the extrenie-llood drainage of Hum- 
boldt River ( via Crescent Valley I : near the valley Hat 
between Carico Lake (usually dry) and the loot ot 
Big .Sage Foothills of the Shoshone R,nige; shown on 
the U. S. G. S. Carico Lake quadrangle { l'-»62) in the 
norlh-eeiilral part of Sec. 16 of T. 2(i N., R. 45 E., near 
the center of Lander County, Nevada (fig. 1 ). Water 
clear; gravel and mud; Mastuiiiiini very dense (had lo 
he removed to catch specimens); 21 C. at 2 p.m. 
Patrick Coffin and D. Erickson, May 25. 1>^)71; UMMZ 
l'>1774 (3 maturing females, 57-64 mm.); dip-net. 

The dace were found only in the spring head. 



about 1 ■ 3 m. m area and 1 dm. deep. The 
outlet stream, which contained much less Nasiur- 
iiiim and was much shallower and only about 0.5 
m. wide, with flow estimated at 0.25-0.,S cubic 
feet per second, flowed into and sank in a gras.sy 
pasture. The local conditions reemphasized the 
ability of speckled dace to maintain a population 
in a miniscule remnant habitat. A few other 
springs mapped in Carico Lake Valley may con- 
tain dace. Mr. Coffin reports that personnel of 
Nevada Department of Fish and Ciame know of 
no other fish in the valley, but that all the springs 
have not been examined. Hall and Iowa creeks, 
tributary to the south end of C arico Lake Valley, 
possibly have retained Rhiiiichlhys. but almost 
surely not trout, as we have previously noted 
( Hubbs and Miller. I'^UXb, p. 36). However, the 
po.ssibility of a former discharge into the Hum- 
boldt River system suggests need for further study. 
Mr. Coffin feels that recent droughts, including 
one ending about 1^)60. might have eliminated 
any fish in the streams. Presumably speckled dace 
are the only native fish in the Carico Lake basin. 
The hydrography of this basin has been treated 
by Everett and Rush ( 1966). 

Location Rl. — liulian Creek (also called C rums 
Creek), a western Inhutarv lo Chestent Valley, just 
above mouth ol cainun near tenabo. near boundary of 
T. 2S-29. in western part ot R. 47 L.; in mideastern 
l_ander County, Nevada, about 5 miles above Dean 
Ranch, near center of valley, about 22 miles south- 
westerly Irom Heow;iwe dig. I). Clear water; sand, 
stones, mud, and trasli; pools and riffles: little vegetation; 
I 7 C. at mid-d:iy (air M). Hubbs family and Miller, 
August 10, 19.^S (M.^S-117); UMMZ 124908 (lh2, 13- 
88 mm.); (i-loot woven-mesh seine. 

This was a small creek (about O.h to nearly 2.0 m. 
wide where fished I ; sub|ecl to Hoods. I he dace here 
were large and conunon. 

We heard at Dean Ranch, which is irrigated by 
the creek water, that the minnows come down 
pipes and ditches from abotit 5 miles above and 
cause trouble at the ranch. We noted that the 
dace were very wary, and tended to rush far 
downstream when disturbed — much more so than 
the spring-inhabiting dace iR. o. rcliijiius) . just 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



107 



previously observed in Grass Valley. Seemingly 
the behavior of this stoek had been conditioned 
by their persistence in a canyon stream. The cir- 
cumstance that this stream prior to ranch use 
had continued onto the valley floor probably led 
to the survival of the dace ( in general, fishes have 
not survived in canyon streams in the Great Basin). 
On our 1938 expedition, we obtained the indica- 
tion by brief exploration and by testimony at Dean 
Ranch that only dace live in Indian Creek and 
that this stream is the only fish-inhabited water 
in the entire valley. Our informer mentioned to 
us that a rancher who had long been in the valley 
had told him years previously that the minnows 
are native and that the waters of the great flood 
about four decades previously ( about 1 900 ) had 
filled not only the sump of Indian Creek just east 
of the ranch but also a series of dry lakes to the 
north, through which the water had overflowed 
on its course into the Humboldt River just above 
Beowawe. where houses had been washed away. 
The pluvial ( and present flood-water ) outlet 
course is detectable from the sump lake to Hum- 
boldt River not only by the chain of dry lakes. 
but also by banks and sand ridges. Since the 
great flood, the water had seldom reached even 
to the dry lakes on the ancient outlet course be- 
tween the sump of Indian Creek and Humboldt 
River. We were told that this sump, locally 
called a "lake." ordinarily fills to a depth of a 
few inches each spring. It retained a little water 
in August. 1938. We were told that trout had 
been stocked in Indian Creek at about the time of 
the great flood, but that they did not survive. 

Location R2. — Bishop Creek, near extreme head- 
waters of Humboldt River, above and helow diversion 
dam. about 2 miles helow Bishop Creek (Metropolis) 
Reservoir, and 2 miles alxive moLith of canyon, in 1 . 3*^ 
N., R. 62E.; Elko County, Nevada. 9 miles (12 by 
road) almost due north of Wells (fig. 12). Rather 
muddy (at high stage: bottom visibility about 0.5 m.); 
very soft mud, sand, gravel, and boulders; generally very 
swift, locally very slight, current; some Poiaiuoi^eion. 
cf. P. pectituiliis. Chara. etc.; 2rC'. above dam. 22" 
below (air 29°). Hubbs family, June 29, 1942 (H42- 
48); UMMZ 141522 (78, 18-60 mm.); 15-foot seine 
with '4 -inch square mesh. 



The dace were not very common below the 
dam, and were rare above. Mature adults were 
taken on a gravel riffle. The stream here was 
partly fed by hot springs, which were said not to 
contain fish. 

Location R,3. — Springs near Town Cacek, near 
main bend of stream, in northern part of T. .^7 N.. K. 
62 E.; Elko County, 0.6 mile west of Wells. Nevada 
(fig. 12). Semi-stagnant; moderately firm peat to ex- 
tremely soft organic mud, with some purple bacteria on 
bottom; virtually no current; marginal rushes, consider- 
able CInirti. and much floating vegetation; 22' C. (air 
26 1. Hubbs family, June 29. ly42 (H42-49); UMMZ 
141524 (164. 15-59 mm.); 6-foot woven-mesh seine; 
many yoimg discarded. 

The dace were moderately common here. A 
local rancher reported that they had formerly 
been very abundant, that they had been used for 
bait, and that they had almost disappeared when 
the ponds had become nearly dry in the drought 
of 1 934. "Swarms of little fish, none of them over 
four inches in length." were reported by Angel 
(1881. p, 18) to occur "in the valley near the 
town of Wells," "in apparently bottomless foun- 
tains of water miles from any surface streams" 
(.see quotation on the flyleaf). 

Population in Ruby Valley. 

Location R4. — isolated spring about 1 km. south of 
the southern boundary of the Ruby Lake National Wild- 
life Refuge in Riil>\ Valley, Nevada, in NW'4 Sec. 7. 
T. 25 N., R. 58 L.; White Pine County. 4.5 miles south 
of Elko County. Nevada dig. 8). Donald E. I^ewis. 
Refuge Manager, August 8, 19fi7 (Sample no. 2); 
UMMZ 186897 (.^2, 14-74 mm.). 

Rclictits was not included in the sample, but 
was obtained about 1 km, to the west. 

During the course of locating extant popula- 
tions of the relict dace, Relictiis solitciriiis. in the 
Ruby Valley National Wildlife Refuge, Mr. Lewis 
surprised us by submitting, along with several 
series of that native fish, this small sample of 
Rhinichrhys osciiliis rohiisliis. After much 
thought, considerable correspondence, and a de- 
tailed study of the specimens, we came to the con- 
clusion that the speckled dace had been Intro- 



108 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



duccd inti) tin.' valley, probably as escaped or 
dLimped bail, and probably from the nearby head- 
waters of Humboldt River. The reasons for so 
thinking were as follows. 

Until sport fishing was vigorously prtMiioted in 
the Refuge, the relict dace was the only fish 
known from the enclosed basin of pluvial Lake 
Franklin, wherein the remnant waters in the area 
of Ruby Lake ( pp. I 98-1 99 ) were being restored 
for wildlife-refuge purposes. 

In 1965 we found that the ponded waters of the 
Refuge were being very extensively fished for 
largemouth bass and other introduced sportfish. 
and that such utilization of the Refuge had been 
promoted by the issue and distribution of informa- 
tiimal ]iosters and leaflets. Ramps were provided 
for the convenience of anglers. The Nevada Fish 
and Game Department was operating a fish 
hatchery within the limits of the Refuge ( wliere 
regulated public shooting was also being pro- 
moted). Because of the remote location, much of 
the angling was by residents of the general area. 
Many of the fishermen came from Wells, the 
nearest town. Fishermen from there and from 
other towns on the Humbiildt River umild have 
had easy access to speckled dace for use as bait. 
As is mentioned above, the minnows (speckled 
dace) of Town Creek, close to Wells, had been 
used for bait. Anglers from the only other towns 
within a reasonable distance, principally Ely, 
would not have had ready access to Rhiiiichlhys. 

The detailed tabulations of characters of the 
varioirs populations of Rhinichlhy.s o.wnlus 
(tables 1 1-1 S) demonstrate in general very close 
agreement between the sample from the L(Kation 
R4 in Ruby Valley and those from the Humboldt 
River headwaters, particularly Town Creek. The 
agreement is complete in the distinctive features, 
noted above (pp. 104-103). of Rhinichlhys 
osculii.\ rohuslii.s. Agreement is excellent in fin- 
ray numbers (table 15). Vertebrae (table 16) 
average somewhat higher than expected, perhaps 
because of the direct action of the environment, 
or because the limited stock of introduced fish 
happened to carry genes for the higher average. 



Agreement is excellent in lateral-line pore de- 
velopment (table 12; fig. 25), but the barbel 
development ( table I 1 ) averages slightly greater 
than expected, perhaps for such a reason as sug- 
gested for the vertebrae. Agreement is adequate 
in other respects. 

It was suspected that the minnows might have 
been stocked for forage or have been distributed 
accidentally long with bass or other gamefish, 
much as l.ucaiiia puivci ( Baird and Girard) was 
established in Utah and southern California 
(Hubbs and Miller, 1965, pp. 48-49), but the 
Refuge Manager at first assured us that this had 
not been done and would have been contrary to 
Refuge policy. (On our urging, the late J. Clark 
Salyer. then head of the federal refuge system, 
issued imiilicit instructions to prohibit introduc- 
tions. ) Later, however, Mr. Lewis obtained from 
the records of the Nevada State Fish and Game 
Department the following account of an earlier 
stocking: 

Scptciiihcr <S. I'JsO: Dace IprL'siimabh Kliiiiich- 
lliy.s (isvuliis ri>hii.\liis] and red-striped shiner 
jobvioiisly Riili(iril.soi!iii\ egrci;iii.s] (No. ."^000) 
were placed in ttie South Samp area (Water 
gape |?|. County Line Pond and Willow Pond). 
Source of fish — Tea Creek. 

Tea Creek location and drainage: lea Creek 
originates in the Jarhridge Mountains. .^0 miles 
north of Deeth. Nevada. Tea Creek flows into 
the Mary's River, then into the HLinihokIt 
River. 

This statement obviously refers to the introduc- 
tion of minnows as forage into the general region 
wherein Rhinichthys was collected. 

After this discussion of the introduction of the 
speckled dace into Ruby Valley from a Humboldt 
River headwater was written, it was noted that 
La Rivers ( 1962, p. 432) had reported an intro- 
ductiof, of Rhiniclulnw osciiliis rohiisln.y into Ruby 
Valley, as follows: 

Ijke all the small "minnows." speckle dace 
have been widely used as bait fish and as such 
have been carried about by fishermen. To 
what e\lent this has occurred is indetermin- 
able, but one instance of official planti[ig of 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



109 



this type is known. In August ot 19.'i|. the 
Nevada Fish and Game Commission obtained 
some specimens from Sadler's Ranch in Dia- 
mond Valley. Eureka County, and planted 
them in Ruby Marsh. Elko County, as forage 
fish for the Largemouth Blackbass fishery 
there. 

Prior to this, the marsh suffered from lack 
of forage fishes, this lack being supplied by 
smaller blackbass, to the general detriment of 
the fishery. The speckle dace seem to be well 
established in the Marsh, hut apparently in 
sections more-or-less inaccessible to blackbass. 

In view of our findings in 1934, the "speckle 
dace" said to have become well established after 
this stocking were native relict dace, rather than 
offspring of Rhinicluhys o.sculiis robiistiis brought 
in from the Sadler Ranch, and it seems probable 
that the stock from there did not survive. Com- 
parison of specimens in 197! showed that the 
series from Bishop Creek in the Humboldt system 
(UMMZ 141522) agreed closely with those from 
Ruby Valley (UMMZ 186897) and contrasted 
with the large collection from Big Shipley Spring 
(at Sadler Ranch) in general appearance, includ- 
ing the strength of the basicaudal spot; in the less 
posteriorly inserted dorsal fin (distance from dor- 
sal insertion to caudal base when stepped forward 
extending in males to some point between front 
of orbit and front part of snout, rather than, usu- 
ally, to some point within or just behind eye, 
and extending in females usually to some point 
within eye, rather than from rear edge of eye to 
well behind eye ) : in the less complete lateral line; 
and, in general, in the slenderer caudal peduncle. 

It will be interesting to determine whether this 
dace will multiply and spread through Ruby Val- 
ley. It apparently would not be prevented by the 
preoccupation of the area by the native relict dace, 
for that species has been so nearly wiped out in 
the valley that it persists only in isolated springs 
to which bass have not gained access (pp. 197- 
199). Very likely this predator will also block 
the establishment of speckled dace. We found no 
evidence that Richardsonius had gained a foot- 
hold in the valley. 

The persistence in the spring in Ruby Valley 



of the characters of R. o. lobiistus has a bearing 
on the genetic integrity of the differentiated pop- 
ulations of other isolated desert springs — con- 
stituting, in effect, a natural experiment (see p. 
80). 

Characters of Populations of Rhinichthys 
osculus robustus sampled from the hum- 
boldt River System. 

Analysis of the characters of these populations 
has shown considerable heterogeneity, though 
within a single general pattern. We briefly treat 
the local variation shown by the samples from 
Carico Lake Valley and Crescent Valley, Bishop 
Creek, and springs near Town Creek; and for the 
stock in Ruby Valley assumed to have originated 
by introduction from a Humboldt River head- 
water. 

Sizi;. The dace of Crescent Valley are among 
the largest of the species, definitely larger than 
tho.se of other Humboldt samples studied (table 
20), which are of average size. They exceed in 
size all other populations of Rhinichlhys treated 
in this report. The large females are roughly 
matched by the larger females of R. o. reliqiiiis. 
but the adult males in Crescent Valley are 
markedly larger than the largest taken in Grass 
Valley. 

Coloration. These series, typical of R. o. 
robiisliis (p. 104), are relatively uniform in color 
pattern and in life color. 

Form. In form of body and head these lots 
are also typical of R. o. robust us. 

Barbel. In four of the five series the barbel 
is more often developed than absent, and is as 
often present as absent in the 3 fish from Carico 
Lake Valley. The barbel in all series is more 
frequently judged "absent" or "small" rather than 
"large" ( table II). 

Lateral line. The development of the lateral 
line, in terms of the number of pores, progresses 
at a relatively similar rate in all Humboldt samples 
(table 12, fig. 25), much as in the races from 
Diamond Valley, but much more rapidly, and 



110 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



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112 



CALIFORNIA ACADEMY OF SCIENCES 



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VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



n3 



much farther, than in the other subspecies here 
treated (figs. 25, 26). The Crescent Valley sam- 
ple, of outstandingly large fish, shows a less rapid 
increase at first, but eventually continues to a 
somewhat higher level. A similar relation holds 
for the Coils Creek sample in the Lake Diamond 
series (p. 104). 

SUPRATEMPORAL CANAL. This canal is some- 
what variably either united or interrupted in these 
five samples ( table 12). 

Morphometry. There are some deviations 
among the four main samples in morphometric 
proportions ( table 13). The dorsal and anal fins 
tend to be more posterior than usual in springs 
near Town Creek. The body and the caudal 
peduncle tend to be deepest in the Ruby Valley 
sample. There is some fluctuation, rather incon- 
sistent, in length of the head and of the head parts, 
but the fin measurements are rather uniform. 

Se.xual dimorphism and nuptial charac- 
ters. The degree of sexual difference in predorsal 
length and in the size of the fins is about average 
and is relatively uniform among the four main 
samples (table 14). On the average, the differ- 
ence in the size of the dorsal fin is greatest in the 
Bishop Creek sample. 

Fin rays. There is little variation among the 
four main samples in number of fin rays (table 
15). For the pelvic count there is a weak cline, 
decreasing eastward. 

Vertebrae. There is also some fluctuation in 
counts of precaudal. caudal, and total vertebrae 
(table 16). The averages are lowest for Bishop 
Creek and highest for the three specimens from 
Carico Lake Valley; the caudal and total counts 
are highest for Ruby Valley (see also p. 108). 
The samples from Crescent Valley and Bishop 
Creek seem slightly but significantly different. In 
the samples from Carico Lake and Crescent val- 
leys the position of the pelvic-fin insertion (table 
17) is farther back on the average, in reference 
to the overlying vertebrae, than in the Humboldt 
headwater series. 

Scale rows. In correlation with the vertebral 
counts the scale-row counts ( tabic 1 8 ) are on the 



high side for the samples from Carico Lake and 
Crescent valleys and on the low side for Bishop 
Creek. 

Gill-rakers. The raker counts are relatively 
uniform. 

Sexual differences in numbers and bio- 
mass. Not .striking (table 19). 

The moderate inter-sample variation within the 
Humboldt River system series is also referred to in 
the following account of the populations in the 
Lake Diamond system. 

Populations of the Pluvial Lake Diamond 
System. 

Field exploration and frequent local inquiry 
have disclosed only five isolated spring popula- 
tions of Rhinichthys oscidiis in the extensive 
drainage system of pluvial Lake Diamond, 
Nevada (pp. 17-20). We think that some addi- 
tional stocks may inhabit springs along an ap- 
parent fault line close to the western edge of the 
playa of the ancient sump lake, particularly in the 
general vicinity of Big Shipley Spring (p. 17). 
The occurrence in Roberts Creek of fish that may 
be speckled dace has been hinted (p. 19). There 
is no evidence that other isolated stocks occur in 
the drainage system. The evidence is further 
discussed following the accounts of the several 
Locations. 

Location R5. — Hot-spring outflow on Potts Ranch 

(previously known as Wilson Ranch), issuing at the 
south end of a lava hill immediately east of Stoneberger 
Creek, a largely intermittent stream, in the Monitor 
Valley division of the Diamond Valley drainage system, 
near the boundary line of Sec. 1-2, T. 14 N., R. 47 E.; 
Nye Coimty, Nevada (fig. !<). Clear water: rather firm 
clay and muck; generally swift; Fotaiiiogclon. cf. P. 
pecrinutiis. over about one-fourth of bottom; 32-34° C^. 
where fished (hotter above — fig. 20: air 24°). Hubbs 
family. August 16. 193S (M38-I32); UMMZ 124*^37 
(103, 19 — KS mm. I; d-foot woven-mesh seine. 

Whether Rhinichllnw still persists in the out- 
flow of the hot spring on Potts Ranch is unknown 
to us. An examination of the spring area (fig. 
20, based on a rough field sketch of August 16, 
1938) suggested that prior to the extensive ditch- 



114 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




Fkilri- 20. Miip of warm springs on Potts Ranch. 
Monitor Valley, N\e County. Nevada I Location k.s). 
sketched, with temperature records addei.1. when tish 
collection M.^,S-122 was made on August Id, 1^)3X: 
thrce-lourths ol the lish were kikcn in the encircled 
area. 

ins: of Ihc duttlow into hay meadows tlic dace 
probably abounded in the cooling stteani courses 
thai iiad previously exi.sted. south of the ranch 
iiouse. in the natural meadow west of the lava 
hill, just as in 19.^8 they still persisted in the 
then still natural outflow of liig Shipley Spring 
(Location R(S). Three-fourths of the dace were 
collected where seepage from the f<irmer meadow 
area below the springs rose in the old channel jiist 
before it received the trenched ditch that bypassed 
the old meadow ( this small area of chief collecting 
is circled on the map; here temperatiites were 
recorded as .^2 and 34 C. ). No fish were taken 
above tliis area, in the trenched i.litch kept clean 



of vegetation, and the only dace seen, at a point 
in the ditch where 37 was recorded, was swim- 
ming erratically against the bank, in Cluira. None 
were seen near the spring sources, in water at 
3X — 14 C, Below the very limited area where 
most of the population was concentrated, in the 
kilometer above the dispersal of the flow in 
ditches, the dace were very few and scattered. It 
seemed c|uite obvious that the handling of the 
water had endangered the isolated, remnant fish 
pc^pulation. It was evident that the supply had 
been decimated to the point where Mr. Potts was 
unsure that any had survived. A few years earlier 
they had been, he reported, very numerous. 

I ocation R5,A. — Hot-spring outflow around I)i;inas 
I'lnHJi l{<)^^l. at source of short flow ol south iork of 
.Sloneherger C feek in Monitor \'alle\ division ol the 
Diamond Valley drainage system, near center of south 
herder of Sec 
Nevada ( fig. .^ ) 
Dianas Punch Howl outflow, ahout 10(1 in. upstream 
from tia\evtine hairier to surface flow; .^7..'^ C: Rohert 



22, T. 14 N„ R.47 E.; Nye County, 
1 hree samples. — ( I ) Labeled as 



L. IJrown, Fehruary 9, 1972 (44, 



-48 mm. 



(2) 1 ahelcd as oLitflow from Dianas Punch liowl, ahout 
2(K) m. downstream from harrier to surface flow; .i7 ' C; 
dissolved o\ygen 2.0 ± 0.2 mg. I.; Rohert L. Brown, 
February 4. 1472 I .s4. 22-4.'i nun.).' (}) Labeled as 
oiittlov\ ol Dianas Punch Bowl, hul as from ahout 
.lOO m. ilow nstrcam from the most western major 
spring; .^4.0 C. ( hv calibrated thermisler recorder): 
dissolved oxygen I..S±0.2 mg./l. ( Hach dissolved- 
oxvgen kit): Rohert E. Brown and Thomas M. Jenkins, 
.Ir.. Fehruary 1 4, 1^)72 (40, .^0-47 mm.).' These spring 
waters are further discussed on p. 20. 

Location \<(t. — Spring-fed pool (ahout .s . 7 m.) in 
grass me;idov\ along the essentially drv Coils Creek, at 
three Bar Ranch. 4}. 5 miles hy Roberts Creek Road 
northwest of luireka I Coils Creek arises between the 
Toivahe Range and the Red HilK section ol Roberts 
Mountains, and flows, in flood, through Koheh V-'alley 
,iikI Bean llal to |Oin the \Uxk\ drainages ol Monitor 
and .Antelope vallevs. thence to pass through Devils 
tjate to enter the drv lake bed ot I5iamontl Valley): 
located near bonndary line T. 22-23 N.. at about midtlle 
of R. 44 L.; near west edge of Ein-eka County, Nevada 
(fig. -^ ) . Moderately clear water: rather firm clay and 
mud; no curient; rushes, lic/iiiscliini. and rufiiiiKiacltm. 

-"■ Ttlc spcciincns so listed .iic Itiose ulili/cd in this papti ; some were 
rcUillKil to the eolleelor, who has t.ikcn other e\.iniples in .l.iniiary 
• iilil 111 .lune. |si7: 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



15 



cf. P. pccriiuiius: 14- C. (air 21 ); Hiibbs family and 
Miller. August 15. 1938 (M38-127); UMMZ 124935- 
36 (1,136. 13-17 mm.): derris. 

Except for a report of fish thought not to be 
trout, which might have been speckled dace, in a 
small valley in the course of Roberts Creek (p. 
19). no indication was obtained of native fish 
life in any other waters in the northern part of 
the western arm of the flood-water drainage basin 
(p. 20). 

Location R7 (and G5). — Two large springs on Birch 
Ranch (Jorge Jacobsen's at time ot collecting; 'Thomp- 
son Ranch" on Ely 1:250.000 map), on east side ot 
Diamond Valley near south end of Alkali Desert, at 
west base of Diamond Mountains, near north margin of 
T. 23 N.. R. 54 E.: close to east boundary of Eureka 
County. Nevada (figs. 3. S). Water (in ditch 2-3 m. 
wide and in pond) clear and somewhat sulphurous: 
fairly firm clay and mud: moderate to slight current: 
considerable growth of rushes. Naici.s iiuiiiiui. U iricidurin. 
etc.: 22'- C. at 7:30 a.m. (air 13 ). Huhbs family and 
Miller. August 13. 1938 (M38-126): UMMZ 124932 
(188, 17-65 mm.): 6-foot woven-mesh seine and 25- 
foot seine with ' 4 -inch square-mesh in bag. 

Mr. .lacobsen, who had been at the Birch 
Ranch for .3 5 years, believed that the dace and the 
chub ( the modified local form of Gila hicolor 
ohi'sd discussed on pp. 18, 154) were native, 
and that they were the only fish on the east side 
of Diamond Valley. We found the two species 
living here together, in both ditch and pond, with- 
out any evident hybridization. No indication was 
obtained that any native fish had survived else- 
where on the east side of Diamond Valley (p. 
154). 

l^ication R8. — Outflow from a small lake (fig. 21 and 
Eakin, 1962, inside front cover), the impoundment of 
liig .Shipley .Spring (recently mapped as Shipley Hot 
Spring), at the Sadler Ranch, south of the ranch houses, 
near middle of west side of Alkali Flat, at the foot of 
Sulphur Springs Range, in T. 14 N., R. 52 I:.: in mid- 
eastern Eureka County. Nevada (figs. 3. 8). Water 
clear, slightly sulphurous: mud, sand, and hot-spring 
deposits: moderate to slight current: considerable rushes. 
Naias marina. Utriciilaria, etc.: 28° C. at 7:30 a.m. 
(air 19"). Hubbs family and Miller. August 13, 1938 
(M38-123): UMMZ 124930-31 (415, 22-55 mm.): 
6-foot woven-mesh seine and 15-foot seine with '4 -inch 




Figure 21. Ponded head of Big Shipley Spring, at 
Sadler Ranch, Diamond Valley. Eureka County, Nevada, 
overlooking Alkali Flat (the bed of pluvial Eake Dia- 
rnond). Outlet of pond (Location R8) yielded a race 
of Rliiniclilliy\ nsciiliis ralyisrus. Photographed by 
Laura C. Hubbs. Auizusl 12. 1938. 



square mesh. The bottom of the pond away from the 
large springs, which issued at 102'^ F. (39° C). at the 
main soince, was covered with V tiimlaiui and Naias 



The dace were found in abundance (and were 
seen nowhere else) in the large outflow stream 
below a natural falls about 2 m. high, especially 
in channels among the rushes. Above the falls, 
and in the large ditch leading toward the ranch 
houses, only goldfish. Cardssiii.s annitus ( Lin- 
naeus), were taken, and only this exotic was seen 
in the pond and in ditches close by; and the gold- 
fish were in all the ditches. The goldfish had been 
planted by the rancher (Mr. Sadler). 

Character.s of Populations Referred 

TO RllINICHrHYS OSCUl.VS ROBUSTUS, IN THE 

Lake Diamond Drainage System. 

Characteristics of the speckled dace from each 
of the five Locations, just described, within the 
large pluvial Lake Diamond drainage basin, are 
different in small ways from one another and from 
those of Rhiuichthys osciiliis robiistiis of the Hum- 
boldt River system. The differences, however, in 
our judgment, are hardly trenchant enough to 
warrant subspecific .separation. The differences 



116 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELIC T FISHES 



117 




Figure 23. Rhinichiliys osctilus rahiisnis. riiciiii from Diamond Valley. Nevada. A. Birch Creek (Location R7); 
UMMZ 124932. no. 21; male. 40.2 mm., side view. B. Same specimen, top view, showing nuptial tubercles and form 
of pectoral fin. C. Big Shipley Spring (R8): UMMZ 124930, unnumbered specimen; female. 53.3 mm. 



Figure 22. Rhiimhthys (KsciiIiis rohiisms. from drainage basins of Humboldt River and pluvial Lake Diamond, 
Nevada. A. Bishop Creek (Location R2): UMMZ 141522.no. 14; male. 44.7 mm. B. Same catalog number, no. 8; 
female, 45.1 mm. C. Coils Creek (R6); UMMZ 124936, no. 29; male, 43.5 mm. D. Same field collection; UMMZ 
124935, no. 2; female, 71 mm. 



118 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



certainly are less impressive than are those that 
distinguish from one another and from R. o. 
robiislKs the populations that we designate below 
by the names R. o. reliqiius. R. o. uliiioponis. and 
R. o. k'thoponis. 

Sizf:. The fish comprising the five populations 
of the Lake Diamond drainage system described 
above differ rather markedly in si/e (table 20). 
but no more strikingly than do the populations of 
the Humboldt River system discussed above (p. 
109). The collections from warm springs (Potts 
Ranch, Dianas Punch Bowl, Birch Ranch, and 
Big Shipley Spring) are, perhaps by direct en- 
viritnmental response, among the most dwarfed 
stocks of Rhinichlhys we have examined. Those 
from Coils Creek are about as large as R. o. 
rclicjiius and. among the populations handled, are 
definitely exceeded in size only by the dace of 
the Crescent Valley population. 

It seems particularly evident that the fish in 
the hot springs about Dianas Punch Bowl are 
dwarfed, and are subject to accelerated develop- 
ment, both seasonally and annually. The sam- 
ples taken in February. 1972. include females 
as small as 29 mm. with ova already enlarging. 
though obviously less than one year old. Some 
in the ."^.5 and 4.0 ..S-cm. groups had nearly full- 
formed ova. Males in the 3.0 to 4.0 groups, in 
part at least surely less than one year old. already 
had very strongly enlarged paired fins, though 
none had yet developed the nuptial tubercles on 
the pectorals. 

Coloration. In general, the coloration cor- 
respontls rather well with that of R. o. rohiiMus 
in the Humboldt River headwaters ( figs. 22. 23 ). 
The primary and secondary bands are apparently 
somewhat more diffuse. The caudal-base wedge is 
somewhat more variable and on the average not 
quite so conspicuous. The blackish stripe on the 
sn(Hit is usually distinct and is subcontinuous with 
the blackened front part of the upper lip. The 
speckling on the body is moderately conspicuous 
m all the series, usually in a rather fine peppery 
pattern. The specimens from Birch Ranch show 
a somewhat greater tendency than the others to 
develop dark horizontal streaks along the scale 



rows on the side of the trunk. In this series, also, 
the general pattern is usually more strongly bi- 
colored than in the others, mainly because the 
lower surfaces are almost entirely silvery on both 
head and body. In these last two respects the Big 
Shipley Spring specimens closely approach those 
from Birch Ranch, with an overlap in variation. 
The specimens from Coils Creek, perhaps as an 
environmental effect, are the darkest, and the 
dark puncticulation tends to extend onto the 
lower side of the head and sometimes to form a 
band across the front of the breast. In all five 
series there is at least a trace of the blackening of 
the crotches of the dorsal and caudal rays, and in 
.some specimens in the forks of the anal rays. 

LiFi-; coiJ)Rs. As noted at the time of collec- 
tion, none of the specimens at Potts Ranch had 
bright colors. Some showed orange-gilt reflec- 
tions over the dark lateral band, and scales, often 
in patches, stood out sharply with silvery reflec- 
tions. 

The fish at Coils Creek were rather brownish- 
olive, with a strong wash of golden tan on the 
mid-sides. The cheeks tended toward gilt, and in 
some the subopercle was almost orange. In many, 
the posterior part of the lateral band was orange- 
gilt. A wide area near the pectoral axil was watery 
orange-tan. The pelvic axil was watery-salmon 
m some, but no such color appeared at base of 
anal, nor about mouth or preopercle. A few had 
a pinkish speck at the upper end of the gill-open- 
ing. Yellowish specks at each end of the dorsal 
base were not conspicuous. Some were orange- 
gilt over almost all of the middle and lower sides. 
One young specimen, 19 mm. in standard length, 
clearly a mutant, was clear pinkish-yellow, with 
no dark pattern; its fins were very pale yellow, and 
its eyes were gilt. 

Birch Creek fish, taken the same day. were 
noted as apparently like those at Big Shipley 
Spring, which were described as distinctly olive- 
green ( perhaps in respon.se to living in submerged 
green-leaved vegetation ) . Blue spangles on scales 
were particularly bright. The lateral band tended 
to be tan. The axils of the paired fins were at 
most very watery-salmon, and there was only a 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



bare trace of this color at base of anal and upper 
end of gill-opening, and none at angle of pre- 
opercie or about mouth. The checks were bluish, 
not gilt. 

Form. The general form in each set of speci- 
mens is rather chubby, probably least so in the 
Big Shipley Spring and Dianas Punch Bowl series. 
In all. the mouth is sufficiently oblique for the 
rostral stripe to include the front of the upper lip. 
The fins are relatively rounded, as is usual in 
forms from springs. 

Barbel. The incidence of barbel develop- 
ment in the five populations in the Lake Dia- 
mond drainage basin fluctuates somewhat, as it 
also does in the Humboldt River system ( table 
11). In three of the five populations, from Potts 
Ranch. Coils Creek, and Big Shipley Spring, 
there seem to be about equal numbers of speci- 
mens with no barbel, with a barbel on one side 
only, and with a barbel on each side. The Birch 
Creek sample, however, includes a high propor- 
tion (69 percent) with no barbel on either side. 
The Dianas Punch Bowl sample shows the 
strongest development of the barbels, thus seem- 
ingly differing from the Potts Ranch sample. 

Lateral line. In all five populations the 
lateral line on the body develops with age well 
toward completion (table 12, fig. 26). The rate 
of pore formation as plotted against standard 
length is least rapid in the least dwarfed popula- 
tion (that of Coils Creek), perhaps because the 
rate is more dependent on age or state develop- 
ment than on size; yet members of this population 
finally develop as many. Similarly, among the 
populations of the Humboldt River system sam- 
pled, the one from Crescent Valley is by far the 
least dwarfed and has the slowest rate of pore 
development in terms of standard length, yet in 
the end attains at least as high numbers. At the 
other Locations in the Diamond system the pore 
development in the dwarfed stocks is more rapid, 
apparently in the close-set sequence of Big 
Shipley Spring, Birch Ranch, Dianas Punch Bowl, 
Potts Ranch. 

Supratemporal canal. The populations 
sharply differ in the proportion of specimens with 



united and interrupted supratemporal canals 
(bottom entry in table 12). In the Potts Ranch 
and Big Shipley Spring samples the canal is usu- 
ally united; in the Coils Creek lot, nearly all are 
interrupted; in the Dianas Punch Bowl and Birch 
Ranch lots, these conditions are about equally 
represented. Similar fluctuations characterize the 
four samples from the Humboldt River system. 

Morphometry. In proportions, there is wide 
variation (table 1.^), with, in general, a broad 
overlap between the five samples measured, as 
well as between them and the samples of R. o. 
mbustiis horn the Humboldt River system. There 
seem to be no very notable differences among the 
samples within the Diamond system or between 
these and the Humboldt basin lots in the propor- 
tions of the head parts or of the fins. 

The feature of the Diamond-system popula- 
tions that early caught our eyes is a more ample 
development than is usual of the trunk region, and 
the compensatory seeming shrinkage of the uro- 
some — features very commonly shown by popula- 
tions living in springs. The morphometric 
analysis, however, does not appear to justify sub- 
specific separation. The better development of 
the trunk region, as expressed by the predorsal 
proportion, indeed seems sharp when the samples 
from Potts Ranch and Big Shipley Spring are 
compared with those from Crescent Valley and 
Bishop Creek, and significantly, when compared 
with the sample of transplants taken in a spring 
in Ruby Valley (further suggesting that the char- 
acter is gene-dependent). However, the samples 
from Coils Creek and Birch Ranch in the Dia- 
mond system are approximately matched in this 
respect by the set from springs along Town Creek, 
a Humboldt headwater, and the specimens from 
Dianas Punch Bowl are intermediate. The con- 
trast between spring and creek populations is con- 
firmed, but subspecies separability is not. 

In the anal-to-caudal proportion there is al- 
most no overlap between the values for either the 
Big Shipley Spring or the Dianas Punch Bowl 
series and the Crescent Valley sample, for either 
sex. but between the other samples there is wide 
overlap or virtually no difference. The distance 



20 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



from caudal base to pelvic insertion when stepped 
forward reaches in the five Diamond-system sam- 
ples measured to any point between front of orbit 
and one eye"s length before tip of snout (with 
some sexual dimorphism), but about the same 
range of variation was noted in samples from the 
Humboldt River system. The position of the 
pelvic-fin insertion, as indicated by the serial num- 
ber of the overlying vertebra (table 17). is es- 
sentially similar among the populations from the 
two drainage systems, with the exception that 
this measure quantifies a statistically more pos- 
terior position of the pelvic fin in the Coils Creek 
sample, as contrasted with all other series re- 
ferred to R. (>. rohiisliis. except those from Carico 
Lake and Crescent valleys. 

The body and caudal-peduncle depths average 
greater in the Diamond system, except for the 
Dianas Punch Bowl series, than in the Hmiiboldt 
basin samples, but the higher values shown by 
the Ruby Valley transplants suggest that these 
features are phenotypic. 

The head averages larger in the Diamond- 
system samples, but the proportions are nearly 
matched by those for the samples from springs 
near Town Creek and from Ruby Valley. 

SFXU.AL dimorphism and NLU'TIM. charac- 
n Rs. In the position of the dorsal fin and in the 
length of each fin, the populations of the Lake 
Diamond system vary widely, with a broad over- 
lap in the values for the po|iulations sampled 
from the Humboldt River system (table 14). 
The Big Shipley Spring sample is remarkable in 
not showing the usual backward location of the 
dorsal fin in females, yet is distinctive in having, 
in males, among all samples analyzed from Ixith 
systems, the greatest relative enlargement of the 
pectoral fin (but not of the pelvic). However, 
the index value is almost as large in the Dianas 
Punch Bowl fish, which, oddly, exhibit stronger 
dimorphism in fin size than those from Potts 
Ranch nearby. Insofar as exhibited, the nuptial 
tuberculation (fig. 2.^B) agrees with that de- 
scribed for the species (p. ^JS ). 

Fin ra^s. For none of the fins are the differ- 
ences in ray number between samples within the 



Diamond system or betv\een these samples and 
the ones for the Humboldt system, as shown in 
table 15, of subspecific significance. There are 
some suggestions of the trend toward a reduced 
number in the spring-inhabiting populations. This 
is hardly show n for the dorsal and anal rays. For 
pectoral rays there is probably a slight numerical 
leduction in the C oils Creek and Dianas Punch 
Bowl samples, as also in the set from springs near 
Town Creek (a Humboldt headwater), and a 
small but seemingly definite reduction in the three 
other samples from the Diamond system. For 
the pelvic rays there seems to be a slight reduction 
for the series from springs near Town Creek, 
oddly for the Ruby Valley transplants, and for 
the Dianas Punch Bowl and Coils Creek samples, 
and a more definite statistical reduction in the 
three other series from the Diamond Valley com- 
plex. 

ViiRTEBRAE. The total vertebral numbers 
( table 1 6 ) show only a very slight and very ir- 
regular trend toward reduction in the populations 
from the Lake Diamond drainage system, and 
some odd, slight fluctuations among the samples 
from the Humboldt River system. 

ScALi-; ROW'S. Within each drainage basin, the 
counts for four scale rows ( table 1 <S ) constitute a 
cline. as follows: 

For the IDiaiiiond basin: (Big Shipley Spring or Birch 
Ranch I • I'otts Ranch • Dianas Punch Bowl 
(except tor rows around peduncle) ■-,: Coils Creek. 

For the Humholdt basin: Bishop Creek < Ruby 
Valley -: springs near town Creek < Crescent 
Valley (with a shift in position for rows around 
caudal peduncle between sprnigs near Town Creek 
and Ruby Valley). 

Between the two extreme samples of the two 
categories ( low coimts for Big Shipley Spring or 
Birch Ranch and high counts for Crescent Val- 
ley ) there is almost no overlap for the predorsal 
series or for rows arotmd caudal peduncle, and 
only a moderate overlap for the lateral-line and 
around-body series. The three specimens from 
Carico Lake Valley had the highest count for 
lateral-line scales, but median values for the other 
series. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



121 



Gill-rakers and pharyngeal teeth. For 
the numbers of gill-rakers (table 18) and teeth 
no notable differences were found between sam- 
ples within the Lake Diamond drainage basin, or 
between those and the series counted from the 
Humboldt River system. 

Sexual differences in numbers, biomass, 
and size (tables 19, 20). In strong contrast to 
the samples from the Humboldt system, those rep- 
resenting the five isolated populations of the Dia- 
mond system vary spectacularly in sex ratio, in 
terms both of numbers and of biomass: the ratio 
of males per 100 females varies from 23 in the 
Big Shipley Spring sample to 124 in the Potts 
Ranch collection. The ratio ( M F ) in terms of 
biomass varies, in essentially the same sequence, 
but to an exaggerated degree, from .12 to 1.10. 

The sexual discrepancy in size also varies. In 
the Potts Ranch, Dianas Punch Bowl, and Birch 
Ranch series, the males and females are both 
dwarfed, and only 30, 6. and 17 percent of the 
females, respectively, are in .5-cm. size groups 
larger than any containing males, suggesting that 
most individuals may be yearlings (the ages of 
the fish were not determined). In the Big Shipley 
Spring sample the males arc about as small as in 
these three sets, but 45 percent of the females are 
in size classes larger than any containing males — 
suggesting a considerable number of a greater 
age. In the Coils Creek series both .sexes are 
larger, indicating better growth, and 1 5 percent 
of the females are in size groups exceeding the 
largest for males. 

Grass Valley Speckled Dace 

Rhinichthys osculus reliquus Hubbs and 

Miller. 

(Figure 24 A. B.) 

Rhinichlhys osculus reliquiis Hubbs and Mm tfr. 1972. 
p. 104 ( diagnosis). 

This subspecies of Rhliiiclithys osculus. one of 
the most distinctive, was, until its almost certain 
recent extermination, confined to Grass Valley, 
the depression that constituted the drainage basin 



of pluvial Lake Gilbert (pp. 10-14, fig. 2), 
near the middle of the northern border of the 
Great Basin, in central Nevada. It apparently was 
the only kind of fish that survived in the basin 
until modern time, though there are reasons (see 
below) to believe that Gila hicolor also occurred 
there at some Pleistocene time. The subspecies is 
known from a single collection, with the follow- 
ing data: 

Location R9. — .Spring-fed creek in a grassy meadow 
in the partly enclosed southwestern arm of Grass Valley, 
13 km. east of Mt. Callaghan. in the course of Callaghan 
(Woodward) Creek, on Grass Valley Ranch, in .SW '4. 
.Sec. 10. T. 21 N.. R. 46 E. (see U. S. G. S. Mount 
Callaghan 15-minute quadrangle): in eastern Lander 
County. Nevada (fig. }). Clear water; soft mud: slight 
to moderate current: generally choked with Naslurtiuni. 
Chara, bulrushes, etc.: 19" C. (air 25"). Hubbs family 
and Miller. .August 9. 19.^8 (M38-116): UMMZ 
12490(1-07 (474. 12-82 mm.): LS-foot seine with '4- 
inch square mesh. 

In 1938 the dace here were common and large. 
Dick McGee, then owner of Grass Valley Ranch, 
said that trout are frequently stocked in springs 
and creeks on the ranch, and we saw one where 
we collected. 

We found no evidence that dace occurred else- 
where in the basin of pluvial Lake Gilbert (p. 
14). 

Inasmuch as a local form of Gila hicolor. as 
well as one of Rhinichllns osculus. occurs in Big 
Smoky Valley, the basin of pluvial Lake Toiyabe, 
which apparently obtained its fish fauna from 
Grass Valley through piracy of a southern tribu- 
tary of that valley (pp. 14-13), it .seems prob- 
able that Gila hicolor formerly existed in the 
basin of Lake Gilbert but failed to survive there. 

The physiographic evidence strongly suggests 
that this relict was derived from the Lahontan 
basin subspecies. Rhinichthys osculus rohustus. 
or its ancestor. That subspecies was found to 
abound in Crescent Valley, into which Lake Gil- 
bert is assumed to have discharged, probably at 
an early pluvial period (pp. 11 and 107), and 
Crescent Valley retains, at high-flood stages, a 
connection with the Humboldt River (p. 107). 



122 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




Fk.lirh 24. Two subspecies ot RlunUlilliys tisculus. from Cirass and Independence \alle\s. Nevada. A. R. o. 
rcliqiiiis. Grass Valley (Location R'J ) : UMMZ 124907. no. 23; parat\ pe. male. 4(i.(l mm. B. R. <>. rfliiiiiiis. same 
field collection; UMMZ 124'-)()(i; holotvpe. female. h7 mm. C. R. <-, U-ilu>iuinis. Independence Valley (R12); L'MMZ 
IX<i51''. no. 13; parat\pe. male. .Ml nini. D. R. o. IciliapiTifi. same lield eoliecliim; L'MMZ IS(i'-)()5; holoI\ pe, fe- 
male, .i5.3 mm. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



123 



However, the highly distinctive characters of R. o. 
reliqiiiis lead us to entertain the thought that it 
might be a relict, which represents some early, 
unknown ancestor (see also p. 14). 

Apparent Extinction. 

When the ditches and the main spring on Grass 
Valley Ranch were examined by the Miller family 
in August, 1 969, in an effort to obtain material of 
this dace for chromosome study, no trace of it 
could be found, despite an all-day effort. Only 
rainbow and brook trout, Salnio gairdnerii and 
Salveliniis fontinalis, were located, and the present 
rancher, Mrs. Knutson (formerly Mrs. Dick Mc- 
Gee), mentioned knowing of large rainbow trout 
below Grass Valley Ranch and of both species on 
and above the ranch. Mrs. Knutson knew of the 
former existence of the dace, and thought that it 
might be extant in a large spring in the meadow 
of Callaghan Ranch, about 10 km. upstream, but 
we found none there. We have since received con- 
firmatory indication of the extinction of this dace 
from Dr. Ira La Rivers (personal comnumication, 
Dec. 9, 1969), who has written us that "I was 
there some ten years ago trying to get the native 
dace and it was gone then, as far as I could deter- 
mine, and the stream was full of trout." He also 
talked with Mrs. Knutson, and added: "i am 
sure the dace are gone from this particular lo- 
cality." Furthermore, a recent effort to locate 
dace in Grass Valley was unsuccessful (Thomas 
P. Lugaski, personal communication, 1972). Ap- 
parently we have described R. o. rcliquus post- 
humously, following an unhappy precedent that 
we established in an earlier publication (Miller 
and Hubbs. 1960, pp. 21-23, 27-28. 56). 

Description and Comparisons. 

Holotype, UMMZ 124906, an adult female 67 
mm. in standard length (fig. 24B). Paratypes 
124907, all other known .specimens (473, 12-82 
mm. long) from same Location ( R9, data given 
above), including the adult male, 46.0 mm. long, 
that is illustrated (fig. 24A). 

The osteological characters attributed to 



Rhiniclithys in the description of the genus 
Relictus (pp. 182-193, figs. 38-45) are based 
primarily on preparations of R. o. reliqiiiis. 

In the following account, R. o. reliqiiiis is com- 
pared primarily with the population of R. o. 
robiistiis that occupies Indian Creek in Crescent 
Valley. The distinctions, however, apply also to 
other populations of that subspecies, including 
those of Bishop Creek and springs near Town 
Creek (pp. 105-121). 

This form is so trenchantly distinctive in com- 
parison with the others herein treated that we 
think it may seem worthy of species rank when a 
sufficient number of other local differentiates has 
been analyzed. 

Size. Although Rhinichthys osculiis rcliquus 
is one of the forms restricted to an isolated spring- 
fed area, it is not dwarfed (table 20), but rather, 
according to our mass collections, reaches the 
largest size (82 mm.) of any of the forms of R. 
osculiis considered in this report, with the excep- 
tion of the population of R. o. robiistiis that oc- 
cupies Indian Creek in Crescent Valley. 

Coloration. In general appearance (figs. 
24A.B) this subspecies is quite different from 
R. o. robiistiis. because the body usually is less 
speckled: the blackened regenerated scales are 
rather fewer and less emphasized and the under- 
lying main dark lateral band is generally broader, 
more solid, and more even-edged. The pattern 
is further intensified by the more definitely 
lightened ground color between this lateral band 
and the dark, broad predorsal stripe. The lower 
sides, especially posteriorly, are often marked with 
deep-lying giant melanophores, which are more 
conspicuous than in R. o. robiistiis and form 
puncticulations somewhat similar to those on this 
region in the subgenus Sipluitclcs of Gila. The 
lower dark lateral band, which is usually rather 
well developed in R. o. robiistiis. and which in 
R. o. Uirivcrsi Lugaski of Big Smoky Valley usu- 
ally is evenly pigmented and strongly deflected 
downward toward or to the anal base, is obsoles- 
cent in R. o. rcliquus. Very distinctive of R. o. 
rcliquus is a dark streak or wedge along the lowei- 



124 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



border of the eaudal peduncle ( it fades in some 
large individuals). The head is characteristically 
darkened from the suborbital region, where the 
dark area is broad, and from the front part of 
the mouth, upward and backward over the front 
and top of the head (whereas in the other forms 
hero treated, the dark area does not extend so far 
below the eye and on the snout tends to form a 
horizontal dark stripe, which is barely even sug- 
gested in R. o. leliqiiHs). The vertical fins also are 
more uniformly darkened, and less speckled, than 
in R. o. i-obiistits, with hardly a trace of the especial 
blackening at the bifurcation of the rays. In this 
respect there is better agreement with the forms 
here called R. o. oligoponis and R. o. Icthoponis. 
In general, however, the color pattern suggests 
that of R. o. robiistiis. 

Another distinctive feature of R. o. reliqiius 
involves the pigmentation of the lower lip: even 
when the lower surface of the head is elsewhere 
devoid of pigment, the lower lip is heavily punc- 
tate ail around. In the populations herein referred 
to R. o. robustiLs\ even in the occasional speci- 
mens that are heavily punctate over the lower 
surface of the head elsewhere, the lower lip, in 
strong contrast, remains immaculate, save oc- 
casionally for a small dash near end of gape, and 
very rarely for partial pigmentatiim aroLuid the 
front rim of the lip. There is a slight tendency 
for the character to break down in the CoWs Creek 
population (referred to R. o. rohii.\lii.s) and in 
R. c>. Icirivcrsi of Big Smoky Valley, and in this 
character R. o. lethoponis ( but not R. o. 
oJiiioporiis) is intermediate. 

Li II-: COLORS. The field records indicate that 
the life colors may also be somewhat distinctive. 
The red color that is often apparent in the axils 
of the paired fins of the species was barely repre- 
sented, in that some individuals had the axils of the 
paired fins and the base of the anal at most rather 
brownish red, seldom at all conspicuous, in fur- 
ther contrast with many populations of Rliinicli- 
tliya osciiliis, there was no red about the mouth 
or preopercle and only occasionally a mere trace 
at the upper end of the gill opening. However, 



the lack of red may have been due to the post- 
nuptial condition of the specimens. The sides were 
silver-spangled on individual scales, but the gen- 
eral tone was brownish olive. The cheeks showed 
gilt and blue reflections in some individuals. The 
field notes further indicate that the general im- 
pression of this dace in life approached that of 
Relictus solitiiriiis. 

Form. Rhinichihys o. reliciuus contrasts with 
R. o. rohustiis in having the body more turgid in 
the nuchal region and in having the muzzle more 
rounded, more declivous, and broader, so that the 
overall width of the mouth approximately equals 
instead of being shorter than the snout. As seen 
from below, the mouth is very broadly U-shaped, 
instead of being narrower, approaching a V. 
These distinctions tend to break down slightly 
when R. o. reliqiiiis is compared with the isolated 
populations of the Diamond Valley drainage, but 
hold well in comparisons with the populations 
from Indian Creek in Crescent Valley and from 
Bishop and Town creeks in the extreme head- 
waters of the Humboldt River drainage basin. 

Frenum. Among 100 specimens of each sex 
specifically examined for this structure, none had 
the rostral groove interrupted by a fleshy bridge 
on the midline; in one specimen the groove was 
shallow. 

Barbi.l (table II). Rhinichihys o. reUqiiiis 
is further distinguished by the virtually invariable 
lack of a barbel. Among 100 specimens of each 
sex carefully checked, only one has a very minute 
barbel, on one side. In all populations here re- 
ferred to R. o. rohiisliis, a considerable proportion 
of the specimens ( a majority for all samples ex- 
cept that from Birch Ranch) have a barbel on 
either one or both sides. The complete or almost 
complete lack of a barbel on either side other- 
wise characterizes R. o. oligoponis and R. o. 
Icthoponis (pp. 100 and 104). 

Latkral-linf, system (table 12, figs. 25, 
26). Despite the fact that it is by no means 
dwarfed, this form, along with R. o. oUgoponis 
and R. o. Icthoponis. is characterized by an ex- 
treme reduction in the development of the lateral 
line on the body. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



125 



The lateral-line system is also degenerate on the 
head. All 40 specimens checked had the supra- 
temporal canal commissure interrupted medially, 
typically very widely. 

Scale structure. The scales show the typi- 
cal structure of Rhiuichthys ( p. 97 ) . 

Morphometry (table L^ ). The morphometric 
measurements provide relatively little that is 
sharply distinctive of this subspecies. The body 
averages slenderer than in other forms. The 
depth of the caudal peduncle is less than that of 
R. o. lethoponis, with only a slight overlap, but 
is about average for R. o. robust us and R. o. 
oligoporus. The head length and the eye length 
are not markedly distinctive, except that they are 
proportionally shorter than in R. o. lethoponis 
(in correlation with the extreme dwarfing of that 
subspecies). The head depth and width and the 
snout length are about average. The interorbital 
width is on the low side. The suborbital width 
is about the same as in most of the other popula- 
tions, but averages greater than in R. o. lethoponis 
( again in correlation with the dwarfing of that 
form). The fin lengths are on the low side, but 
only on the average, and, as is usual in spring- 
inhabiting minnows, the fins are rounded. 

The position of the fins ( table 1 3 ) provides 
some distinctions. The predorsal length is high 
on the average, but variable: in the adult males 
about the same as in the Big Shipley Spring pop- 
ulation, but somewhat greater than in the others: 
in the females, close to the means of all other 
populations sampled. The distance from anal 
origin to caudal base is definitely lowest in each 
sex, but with wide overlap. Very distinctively, the 
pelvic fin is more posteriorly inserted in R. o. 
rcliqims than in any other form here considered 
(including R. o. lariversi of Big Smoky Val- 
ley). This distinction can be shown in two ways: 
( 1 ) the distance when measured forward extends, 
with little variation, about to middle of eye in 
R. o. reliqiiiis, but about to tip of snout in the 
other forms, with considerable variation; (2) the 
pelvic insertion, as seen on radiographs, usually 
lies in R. o. reliqiiiis below a more posterior verte- 
bra (table 17), but this measure of the position 



seems to be less consistent than is indicated by the 
first method and is somewhat subjective. 

Sexual dimorphism and nuptial charac- 
ters. The sexual dimorphism in the length of the 
pectoral fin in this subspecies is more extreme 
than in any of the other populations treated 
(table 14), more because of shortness in females 
than of extra length in males: on the average the 
fin is 6.7 percent of the standard length longer in 
males than in females. The extreme excess length 
in males does not hold for the pelvic fin. The 
dorsal and caudal fins on the average are larger, 
but not quite so extreme. The dorsal fin is, on 
the average, even more posteriorly inserted in 
males than in females, contrary to the situation 
in the species as a whole, and in cyprinids in gen- 
eral. 

The pectoral fin in males has very thick and 
firm rays. The adult males, taken probably past 
the peak of nuptial activity, have, as is usual in 
the species, the outermost, unbranched pectoral 
ray and the two or three following branched rays 
strongly dilated and more or less curved. The 
first ray .seems to have been considerably padded, 
but in the specimens at hand is devoid of nuptial 
tubercles. The tubercles, one per segment, seem 
to have been confined mostly or entirely to the 
first few strengthened branched rays. It could 
not be ascertained definitely whether the single 
row of tubercles branches once, as it often does 
in R. osciiliis. 

Fin rays ( table 13). The counts of dorsal 
and anal rays average very slightly higher than in 
the other populations: contrary to the general 
rule for R. osciilus, counts above 8 in the dorsal 
fin and above 7 in the anal fin exceed those below 
these numbers. The pectoral-ray counts average 
fewer than in any of the populations of R. o. 
robustiis treated (except for the set of 3 specimens 
from Carico Lake Valley), but about the same as 
in R. o. oligoporus and R. o. lethoporus. The 
pelvic-ray numbers average slightly lower than 
in R. o. robust us. but barely less than in some of 
the isolated colonies in the Diamond Valley drain- 
age basin, and very slightly higher than in R. o. 
oligoporus and R. o. lethoporus. 



126 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



80 



70 

CO 

LlI 

cr 
o 

Ci- 60 
LlJ 



< 

LiJ 
t- 

< 



O 



< 

LlI 



50 



40 



UJ 50 

DQ 



20 



10 







^^SCENT VAL 



P^ 



/ 



Springs near Town Creek 
(symbols omitted for 84 

specimens 15-21 mm long, 

all with 0-0 pores ). 



e/ 

O 



LEV 



V 



2 




oligoporus 

j 9.5mi S^ of WELLS 




a 



f 



h/ oligoporus 

,"18 WARM SPRING, CLOVER VALLEY 



- lethoporus 

INDEPENDENCE VALLEY 



20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 



STANDARD 



E N G T H mm 



FiGiM^E 25. Development of lateral-line pores on body in subspeeies of RhiiuclilJiys nsciiliis from headwaters of 
Huniholdt River and from hasin of pluvial Lake Clover, Nevada. Numerals beside symbols indicate numbers of sides 
eiHinted (omittei.1 on entries for specimens less than .^2 mm. lony ha\ing porc-niimber average less than 2.0). Data 
from table 12. 



Vertebral (tabic 16). The piecaiidal verte- 
brae average nioclerately to very sligittly higher 
than in the populations referred to R. o. robustus 
(the Carico Lake Valley set of three specimens 
e.xcepted) and R. <>. Icllioponis. but not higher than 



in R. (>. <'//,t,'()/'<"'"v. The caudal vertetirae. iiow- 
ever. dcfiintely average fewer than in the other 
forms ( though iiardly different from ime jiopula- 
tion of A'. ('. olli^cponis) ; this is surprising, in 
view of the tnore posterior position of the anal 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



127 



(I) 

LlJ 

cr 
o 

CL 
UJ 



< 
or 

UJ 

H 

< 

_l 

Li_ 

o 
cr 

UJ 



< 

LxJ 



70 



60 



50 



40 



30 



20 



- 







15 




STANDARD LEN GTH, mm 



Figure 26. Development of lateral-line pores on body in populations of Rliinu lilliys i>.s( iihi.s rohustus from basin 
of pluvial Lake Diamond, and in R. o. reliquus of Grass Valley (basin of Lake Gilbert), Nevada. The crosses (not 
interconnected) represent the values for Birch Ranch. Numerals beside symbols indicate number of sides coimted 
(omitted on entries for specimens less than 24 mm. long having pore-number average less than l.(J). The values for 
Dianas Punch Bowl Springs, not plotted, appro.ximate those for Potts Ranch or Big Shipley Spring. Data from 
table 12. 



and especially of the pelvic fin in R. o. reliquus. 
Because the higher average for precaudal counts 
is less than the lower average of caudals, the total 
number averages barely lower in R. o. reliquus 
than in some populations of the other subspecies. 



Scale rows (table 18). The four counts, 
namely lateral-line series, predorsal rows, around 
body, and around caudal peduncle, average con- 
sistently higher than in the two dwarfed sub- 
species [R. (>. olii^onorus and R. o. lethoporus). 



128 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Predorsal 

length 


Dorsal 
fin 


Caudal 
fin 


Pectoral 
fin 


Pelvic 
fin 


-13 
-19 
-14 


8 

24 

8 


15 
14 
10 


39 
46 
43 


20 
25 
16 



1 



21 



28 
26 



If 



14 



19 



52' 
21 



48 



Tahie 14. ScAiiiil (liiiuirpliism in prednrsal ami fin lenulhs m /ui/niUiiions nf Rhinichthys osculus in certain basins 
in Nevada, expressed as excess for males over females in mean values in permillage of standard length.^ 

Subspecies 

Pluvial lake system 
Locality 

R. o. rohiiMi{s 

L. Lahonlan (Humboldt R. ) 

Crescent Valley (10/10)" 

Bishop Creek (7/12) 

Springs near Town Cr. (20/20) 
L. Franklin (introduced) 

Ruby Valley (4/7) 
L. Diamond 

Polls Ranch ( 10/101 

Dianas Punch Bowl (10/10) 

Coils Creek (20/20) 

Birch Ranch (20/20) 

Big Shipley Spr. (15/21) 
R. o. rcliquiis 
L. Gilbert 

Grass Valley (20/20) 
R. o. oligoporirK 
L. Clover 

9.5 mi. S. of Wells ( .V4 ) 

Warm Sprs., Clover V. (12/15) 
R. (). Iclhoponis 
L, Clover 

Independence Valley (12/12) 



-19 

-10 
-10 

- 7 

-17 

1 



4 
-18 






(-11)^^ 


38 


21 


12 


55 


14 


10 


52 


16 


20 


44 


22 


19 


57 



67 



53 
43 



52 



26 

9 

21 
19 

22 
12 



16 



26 

28 



1 Based on valueb in table l.^. 

" Figures in parenllieses indicate tlic number of adull males and adult females, respectively, that were measured; a lew fin measurements were 
missed. 

"Only ,1 measurements for males and only 1 for females. 
' Only 2 measurements for males and only 1 for females. 



witlt relatively little overlap. Almo.st all averages 
are also higher than in the popiilation.s of R. o. 
rohiislus stucJied, except for the collections from 
Crescent Valley. 

Gn.i.-RAKi.Rs (table IS). The rakers may 
average slightly lower in ntimber than in the other 
populations here analyzed, except for those com- 
prising the type series of R. o. oVigoponts and for 
the set of three specimens from Carico Lake Val- 
ley. 

PnAR^N(;l AL Ti-iTH. In all 20 specimens ex- 
amined the tooth formula was interpreted as 1, 
4 — \. I, by including in the count what appear 
to be alveoli of lost teeth, especially in the lesser 
row (only one such alveolus was largely filled in 
by bone). 

SE.XUAL DIFFERENCES IN NUMBERS AND BIO- 
MASS (tables 19, 20). The type collection of R. 
o. rclii/niis, the only one obtained, comprises, 
young included, 183 males 12 to 47 mm. in 



standard length and 291 females 17 to 82 mm. 
long, a ratio of 63 males to 100 females. The 
discrepancy by volume is even greater, 136 to 
498 ml., a ratio of .27:1. Since the sample was 
collected with a seine having a '4 -inch square 
mesh, most of the young, which were presumably 
abundant on August 9, no doubt escaped. Fe- 
males of the age-group were mostly in the 2.0- 
and 2.5-cm. size-classes, which graded into the 
adult size-classes. Since the adult males were 
small, the young males did not form a distinct 
mode. The size-class fret|uencies suggest that the 
adult males were almost all yearlings, whereas the 
skew size distribution for females, with modes at 
the 3.5-4.0 and at the 6.0-6.5 cm. size-classes, 
suggests that a considerable proportion of the 
females were two years old. 

Di;rivation of Name. The meaning of 
reliifim.s is given as "that is left behind, or re- 
mains." 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



129 



Clover Valley Speckled Dace 

Rhinichthys osculus oligoporus Hubbs and 

Miller. 

(Figure 27.) 

Rhinichthys osculus oligoporus Hubbs and Miller, 
1972, p. 105 (diagnosis). 

This is one of the two spring-inhabiting sub- 
species of Rhinichthys osculus that we recognize 
from the basin of pluvial Lake Clover, which oc- 
cupied the broadly connected Clover and Inde- 
pendence valleys, in northeastern Nevada (pp. 
29-32; figs. 8. 1 1, 12). The second form is R. o. 
lethoporus. described below, from a spring in 
Independence Valley. 

We refer to R. o. oligoporus two slightly dif- 
ferentiated forms, from separate springs, both in 
Clover Valley, near the western edge of the 
ancient lake bed. These spring waters were 
treated by Eakin and Maxey (1951b), with chemi- 
cal analyses of Warm Springs. 

Location RIO. — Spring pond and outflow 9.5 miles 
south of Wells, issuing from near an outcrop of lava 
at the soutlieastern base of a juniper-covered hill, near 
the north end of the western border of the flat bottom 
of Clover Valley, near corner of Sees. 1 9-20 and 29-30. 
T. 36 N.. R. 62 E. (see Elko 1:250,000 map): east of 
center of Elko County, Nevada ( fig. 12). Two collec- 
tions were made, under different conditions: 

First collection, in spring-head reservoir about 3 m. 
deep and about 1 acre in extent, and in the outlet, on 
what was then known as Ralph Ranch. At that time the 
pond was bordered on one side by a wooded slope and 
on the other side by a grassy meadow. Clear water: 
rather firm, whitish bottom: current moderate in outlet; 
choked with Potainogetoji, cf. P. pcctiiuuns, Cluini. etc.; 
water cool. Hubbs family, September 14, 1934 (M34- 
213): UMMZ 132192-93 (94, 14-40 mm.); 6-foot 
woven-mesh seine and 15-foot seine with '4 -inch square 
mesh. On this occasion, in the year of excessive drought, 
this spring, the largest in the valley north of ''Clover 
Ranch " (now Warm Springs Ranch, location Rl 1 ), was 
the only source of water at this major ranch, since 
snow water from the East Humboldt Range had failed, 
and the extensive meadows had "burned up." 

Second collection, in the southernmost of the ditched 
outlets, with one haul in the headwater reservoir, now 
largely silted-in and surrounded by dense woods, on the 
property now called Wright Ranch. Water clear, but 



very easily muddied; bottom now of deep mud; dense 
growth of Potamogeton, cf. P. pectinalus. and a sparse 
growth of a floating filamentous green alga; 17.2-17.8° 
C. ( air 24" ) . Miller family and the Hubbses. August 26, 
1965 (M65-35); UMMZ IS652I (102, 14-55 mm., 
including the female figured); 12-foot woven-mesh seine. 

When these spring waters were examined in 
1934, the dace were scarce and mostly very small, 
apparently because they had been preyed upon by 
large rainbow trout, Salmo gaiidnerii (one seen), 
and because the water had been used for irriga- 
tion of the hay meadows. Diligent collecting pro- 
duced only young fi.sh and a single adult. The 
rancher ( Mr. Ralph ) informed us that previously 
the minnows had been more abundant and had 
reached a larger size. In 1965, a year of heavy 
precipitation, water was more profuse, and trout 
were not in evidence. The population of dace 
had recovered, and adults were not so extremely 
scarce. 

Location R 1 1 . — Warm Sjnings, Clover Valley, at 
Warm Creek (formerly Clover) Ranch, near south- 
eastern corner of Clover Valley, near foot of bajada 
just above the flat ancient bed of Lake Clover, in Sec. 
7. T. 33 N., R. 61 E. (see Elko 1:250,000 map): in 
southeastern Elko County, Nevada, at east side of U. S. 
Highway 93, 3.1 miles north of junction with State 
Highway 1 1 (figs. 8, 12). The main spring was said by 
the owner of the ranch, in 1965. to deliver 3.200 gallons 
per minute. Two collections were made; 

First collection, in the outlet of the deep spring-fed 
reservoir. Water clear; gravel, mud, and debris; vegeta- 
tion dense; 18.5° C: width to 3 m. and depth to 0.6 m. 
James E. Deacon and Mary Beth Rheuben, September 
14, 1964 (JED 64-55): UMMZ 186902 (holotype, 55.2 
mm.) and 186903 (29 paratypes, 15-58.5 mm.); 15- 
foot seine with '4 -inch square mesh. 

Second collection, in a very short lateral inlet flow, 
0.3 to 1 .0 m. wide, of the reservoir. Water very clear, 
but easily muddied; silt over gravel, rocks, and firm 
mud; current barely perceptible: largely choked with 
Nasturiiuiii. with some fine-leaved submergent, and 
some Cham toward spring head; 19.3° C. (air 25"). 
Miller family and the Hubbses. August 26, 1965 (M65- 
34); UMMZ 186520 (88. 12-28 mm.); 6-foot woven- 
mesh seine. 

When this spring, then known as Clover Spring. 
was first examined by us, on September 14, 1934. 



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MEMOIRS 




Figure 27. Rhinidilhv.s dsculiis ulii^oporus. from Clover Valle\, Nevada. A. South of Wells >^). 5 miles (Location 
RIO, first collection); UMMZ 132192, unnumbered specimen; male, 39.9 mm. B. Same Location (second collection): 
UMMZ 1 8652 1, no. I; female, 55.4 mm. C. Warm Springs (Rll, first collection); UMMZ 186903, no. 18; para- 
type, male, 44.2 mm. D. Same field collection; UMMZ 186902; holot\pe, female, 55.2 mm. 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



131 



the rancher said that it has a temperature of 
55° F. ( 13° C.) and is fishless, but he may have 
thought of game fish and was, almost uniquely 
in our Great Basin experience, definitely in- 
hospitable and uncooperative. Although we saw 
no fish in this spring, the possibility seems to us 
rather remote that the dace were actually absent 
in 1934. One reason for so thinking is that the 
only reasonable source of dace for stocking 
would have been from Ralph Ranch ( Location 
RIO), which is on the same road, the only one 
on the western side of the valley, or in an im- 
mediately adjacent locality, and the two stocks 
seem to be somewhat differentiated. Further- 
more, the dace may have been in seclusion in the 
ponded spring water, just as Relictiis solitarius 
apparently was on the same day near Currie, fol- 
lowing a night when water froze in camp (p. 
204). 

When the first collection was made, just 30 
years to the day after our first visit to the spring. 
Dr. Deacon observed a sizable population in the 
deep reservoir, and the rancher said that the min- 
nows abounded in the pond. He refrained from 
collecting there because the rancher did not want 
him to disturb the rainbow trout, which had been 
stocked recently, and had reputedly doubled in 
size over the summer. For this reason, the col- 
lecting was confined to the outlet ditch, where the 
dace were scarce, as also in the distributaries in 
the meadow. When we examined the spring about 
a year later, making the second collection, we 
saw no trace of dace in the clear water of the 
reservoir, in the main spring inlet issuing (at side 
of highway ) from just above the bajada base, in 
the main outlet ditch from the impoundment, or in 
the smaller springs and overflows on the meadow 
below. The only place where fish were found, 
after prolonged search, was in a slight lateral in- 
flow, tributary to the main inlet between the 
spring source and the reservoir, and here only 
young fish were encountered. It appeared that 
the introduced rainbow trout were forcing the 
localized form of speckled dace definitely into 
the seriously endangered category. Bullfrogs, 



Rcma catesbeicina, found in 1965 to be common, 
may have contributed to the decimation of the 
dace. 

We assume that this local form, as well as R. o. 
lethoponis of Warm Springs in Independence Val- 
ley, the next subspecies, was derived from the 
stock of /?. o. robustits, or its ancestor, in the 
headwaters of the Humboldt River immediately to 
the north. The indication has already been pre- 
sented (pp. 30-31) of a presumably Pleisto- 
cene, but not latest Pleistocene age, outflow from 
pluvial Lake Clover into the Humboldt River 
headwaters. The marked differentiation of these 
two subspecies from R. o. robiistiis of the upper 
Humboldt River system and the slight differentia- 
tion of the two from one another ( as specified in 
the account of R. o. lethoponis), and the slight 
distinction between the two populations of Clover 
Valley (Locations RIO and Rll), specified be- 
low, are compatible with this physiographic evi- 
dence. 

Warm Springs in Independence Valley have 
presumably long been isolated from the habitats 
of R. o. oligoporiis. and neither a field reconnais- 
sance nor a scrutiny of the Elko 1 :250.000 map 
has yielded any plausible present-time floodwater 
connection, although the flood channel from Lo- 
cation RIO for R. o. oligoporiis runs into the 
alkali flat of Independence Valley. The channels 
in Clover Valley, however, appear to be braided, 
and it is possible that at times flood water dis- 
charges from Location RIO have reached either 
the impounded waters or a flood overflow of 
ephemeral Snow Water Lake, which is fed by the 
discharge of the Warm Springs of Clover Valley 
(Rll). No such ephemeral lake exists in In- 
dependence Valley. Hence it is plausible to as- 
sume that the two populations treated as one sub- 
species ( oligoporiis ) have been connected after 
the Independence Valley form ( now modified as 
R. o. lethoponis) was separated, but it seems im- 
probable that such connection has been recent. 
(In interpreting the 1:250,000 maps involved — 
Elko and Wells — it needs to be kept in mind that 
the 5,600-foot contour in Independence Valley 



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CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



and the 5,700- and 3.800-foot contours in both 
valleys were inadvertently not marked as en- 
closed.) 

Description and Comparisons. 

Holotype, UMMZ 186902, an adult female 
55.2 mm. in standard length (fig. 27D). Para- 
types. 186903 (29, 15-58.5 mm. long). The 
specimens designated as types comprise all taken 
in only the first of the two collections described 
above for Location Rll, including the mature 
male, no. 18, 44.2 mm. long, that is illustrated 
(fig, 27C). 

We c(Miiparc this subspecies, one of the spring- 
inhabiting forms of R. osciilus. primarily with the 
upper Humboldt stocks of R. o. robiistus. Rliinicli- 
tliy.s (). rcliqiiiis is distinguished above from this 
form, and the distinctness of R. o. Icllioponis is 
detailed in the next sub.species account. R. o. 
ollgoponis seems to be much less sharply differ- 
entiated than R. o. rcliqinis. and somewhat less 
modified than R. o. Icllioponis (of the same 
basin). 

Sizi-. Although relatively few adults were ob- 
tained, this subspecies appears to attain about 
average size for R. osciiliis ( table 20 ) . 

CoiORAiTON. In comparison with R. o. rohiis- 
liis ( figs. 22, 23 ), the body in this subspecies ( fig. 
27 ) is more extensively .speckled with black. The 
lower lateral band as a rule is much less de- 
veloped, or not evident; the dark pigmentation 
around the snout is more generally diffused, with 
virtually no evidence of the usual horizontal 
black streak in front of the eye, but retaining a 
tendency for its continuation across the opercle 
that is characteristic of R. o. rohiislKs. The jet- 
black wedge that is ordinarily developed at the 
base of the caudal on the midline in R. o. 
robiistus is much smaller, less intense, and more 
disrupted, and occasionally hardly evident. Dusky 
dashes on the doisal and caudal fins lend to be 
more numerous than in R. o. robiistus, but are 
virtually unrepresented on the anal fin, where 
they are rather often developed in R. o. robiistus. 
These marks are much less apt to form in. and to 



be largely restricted to, the crotches of the bifur- 
cating rays. The preserved specimens as well as 
individuals in life (.see below) have a light streak, 
bordering the main lateral band above, that is 
hardly evident in R. o. robiistus. Despite these 
various differences, the general coloration of the 
two subspecies has many features in common. 
Many of these differences tend to break down 
somewhat when comparison is made with the 
five populations from isolated parts of the Dia- 
mond Valley drainage system. 

LiFF COLORS. The only field notes on color, 
taken at the first collection at Location RIO, 
indicate that the population there is bright olive 
or golden green on the back and silvery below, 
with an intervening bright gilt stripe and with 
dusky mottling. The only adult taken here, a 
male, unlike the specimens of R. o. rclicpiiis, dis- 
played clear red in the axils of the paired fins and 
along the base of the anal. 

Form. In comparison with R. o. robiistus (figs. 
22, 23 ), R. o. oliiioporiis (fig. 27 ) differs in gen- 
eral form. The outlines of the body, and especially 
of the head, are more curved, and the head in par- 
ticular is more rounded, in both dorsal and 
lateral aspects. The mouth tends to be more 
definitely lower than the lower border of the eye, 
and is generally more curved. The whole aspect 
is bulkier. 

Fri-num. Not one of the specimens was ob- 
.served to have a frenum. 

Barbf.l (table II). In this subspecies, like 
R. o. rcliqiius and R. o. Ictlioporiis. the obsoles- 
cence of the barbel that is often evident in iso- 
lated stocks and derivatives of R. o. robiistus has 
been carried almost if not completely to the de- 
gree of total absence. Not one of the 51 speci- 
mens, about equally from both Locations, that 
were checked, on both sides, shows any trace of 
this sensory structure. 

Lati:ral-line system (table 12;figs. 25, 26). 
Perhaps in correlation with its isolation, this sub- 
species, along with R. o. reliquiis and R. o. Ictlio- 
poriis. is characterized by an extreme reduction of 
the lateral line on the body. In this respect the 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



133 



two populations referred to this subspecies are not 
markedly different. 

The lateral-line system is also degenerate on the 
head. The numbers of specimens with a united 
and an interrupted supratemporal canal ( table 
12), respectively, are 3 and 3 for Location RIO. 
and 14 and 1 for Rl 1, suggesting some local dif- 
ferentiation. 

Scale structure. The scales show the typi- 
cal structure of Rhinichthys (p. 91). 

Morphometry (table 13). The morphometry 
of R. o. oligoponis is not sharply distinctive. The 
specimens from Location RIO are among the 
deeper-bodied of those from all the collections 
considered, whereas those from R I I are among 
the slenderer. This difference may be phenotypic. 
Caudal-peduncle depth is about average. Per- 
haps because of the small sample, or because of 
differential nutrition, the three adult females from 
Location RIO have the head deeper and wider 
than do the four adult males from the same place, 
or in the adults of either sex from R 1 1 . The sub- 
orbital seems to average slightly narrower than 
in other populations of the species examined, R. 
o. k'thoponis excepted. Otherwise, the dimensions 
of the head parts seem to be about average. The 
size of the fins is also abiiut average. As in the 
other forms under discussion, the fins are rela- 
tively small and rounded. 

The position of the dorsal and anal fins ( table 
13) is not distinctive among the forms of 
Rliinichthys osciiliis here treated, but the pelvic- 
fin insertion averages farther back than in typical 
R. o. lohustiis or in R. o. lethoporiis, but farther 
forward than in R. o. rcliqims. In this subspecies, 
the distance from base of caudal to pelvic-fin in- 
sertion, when stepped forward, extends to any 
point one full eye length behind, to slightly before, 
rarely to as much as one eye length before, tip of 
snout: in typical R. o. robuslus. to any point one 
pupil length behind to more than an eye length 
before tip of snout ( in populations in springs of 
the Diamond Valley drainage the position of the 
pelvic may be intermediate). In radiographs, the 
pelvic insertion (table 17) lies below the 14th 



to 16th, usually the 15th or 16th, vertebra in R. 
o. oligoponis; usually below the 14th to 1.5th in 
typical R. a. robiistiis; usually below the I 5th and 
1 6th in some populations in the Diamond Valley 
drainage; usually below the 16th or 17th in R. o. 
reUqiius; and below the 15th in R. o. lethoporiis. 

Sexual dimorphism and nuptial charac- 
ters. Sexual dimorphism in the size of the fins 
( table 1 4 ) seems to be about average ( the exces- 
sively large difference in the size of the caudal 
fin and the similar values for the sexes in the 
position of the dorsal fin in the fish from Loca- 
tion RIO seem attributable to the very small 
sample of adults). 

The nuptial tuberculation agrees with the ac- 
count given under the species heading (p. 98). 

Fin rays ( table 15). The counts of dorsal and 
anal rays are usually 8 and 7, respectively, as is 
characteristic of Rhinichthys oscuhis in general, 
but the rays in the paired fins are somewhat re- 
duced in average number: this subspecies and 
R. o. lethoporiis are the only ones among those 
under consideration that seem to have 6 pelvic 
rays oftener than 8, and these two and R. o. 
reliqiiiis. and the set of three specimens of R. o. 
rohiistiis from Carico Lake Valley, are the only 
ones that have 12 pectoral rays oftener than 14 
or 15. The principal caudal rays are normally 
19, of course, but many vary more than usual, 
from 1 8 to 2 1 . 

Vertebrae (table 16). The total vertebral 
counts are about as usual, but the precaudals 
seem to average slightly higher and the caudals 
slightly lower than usual. 

Scale rows. The four sets of scale counts 
(table 18) average definitely lower than in R. o. 
reliqiiiis. about the same or not quite so low as in 
R. o. lethoporiis. and somewhat lower than in R. 
o. rohiistiis ( except for some of the counts of the 
isolated populations of the Diamond Valley drain- 
age system ) . 

Gill-rakers. The rakers (table 18) at Loca- 
tion Rl 1 .seem to average very slightly lower than 
in other forms, but at Location RIO on the high 
side. 



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MEMOIRS 



Pharyngeal teeth. The \5 specimens ex- 
amined for dentition yielded more than usual 
variation; 13 have the typical formula 1, 4 — 4, 
1, but one each has (), 4 — 1. 1 ; 1, 4 — 4, 0; and 
1. 5 — \. 1. The two listed as having no tooth in 
the lesser row of one side show no trace of an 
alveolus there. The count of 5 teeth on the main 
row of the left side of one apparently quite nor- 
mal specimen, is. so far as we know, the only 
record of 5 for the entire genus. More than usual 
variability is suggested. 

Si:XUAL DIFFERENCES IN NUMBERS AND BIO- 
MASS (tables 19, 20). The depleted populations 
of R. (>. (>lii^oi>oriis sampled contained few adults. 
The two collections from Location RIO, taken 
in August and September, contained 189 young 
14 to 30 mm. in standard length, 3 adult males 
40 to 41 mm. long, and 4 adult females 51 to 55 
mm. long. Assuming thai the un.sexed young 
were equally divided by .sex in numbers and size, 
the sex ratio is essentially equal, and the volumes 
are IX. 5 ml. for males and 28.5 ml. for females, 
a ratio of 0.65:1. 

The two collections from Location Rl I, also 
taken in August and September, ctMitained 90 
young 12 to 28 mm. long, 12 adiilt males 28 to 
44 mm. long, and 16 adult females 28 to 59 mm. 
long. Assuming that the unsexed young were 
equally divided by .sex in numbers and size, the 
sex ratio is 93 males to 100 females, and the 
volumes are 17.6 ml. for males and 43.^) ml. for 
females, a ratio of 0.40:1. 

DiRiVATiON OF NAME. The name oliiidi'oni.s 
was derived from I'.A/yos-. few, and tt.^jos, a passage 
through the skin. It is regarded as in the adjectival 
form. 

Independence Valley Speckled Dace 

Rhinichthys osculus lethoporus Hubbs and 

Miller. 

(Figure 24C.D. ) 

Rhinii lilhv\ asciilns Ictluiporus Hlisiss and Miller, 
l')72. p. 105 (diagnosis). 

This subspecies, along with Gila hicolor isohita 
{ p. 32, fig. 12), is confined to Warm Spiings, the 



single spring complex in the entire expanse of 
Independence Valley (part of the basin of pluvial 
Lake Clover), in east-central Elko County, 
Nevada. These springs were mapped and briefly 
described, as "Ralph's Warm Springs," by Eakin 
and Maxey (1951b). The very limited other 
waters in the valley were definitely indicated by 
field reconnaissance to be fishless (pp. 31—32). 
The hydrographic and speciational relationship of 
this subspecies to the other subspecies occupying 
the basin of pluvial Lake C4over are discussed 
above under the heading of that form, R. o. 
olii^oponis. 

Location K12 (and Cill). — Warm Springs of In- 
df])ciitk'ncf Vallev, a spri[ig complex on the west 
margin ol the north arm ol the valley, just oft the base 
ot Pequop Moimtains, approximately on the edge of the 
bed of pluvial Lake Clover, just below the 5,7()()-foot 
( 1,737-m. ) contour (on Elko 1 :250,000 map), on either 
side of the T. 35-.^(i line near middle of K. hd E., in 
east-central Elko County, Nevada (fig. 12). Water very 
clear, but easily muddied; thick clay-mud, firm in places; 
dense vegetation over bottom, very largely Chord, with 
some Myi'ioi'hylliiin. C'lKiii'p/iylhiiii. a broad-leaved 
Potaimiiicton, and a sparse green alga; 26'^ C. (air 21"). 
Miller and Hubbs, .August 25. 1465 (M65-33); UMMZ 
186519 and 1X6905 ( 101, IS-39 mm.); 12-foot woven- 
mesh seine and i 5-foot seine with '4 -inch square mesh. 

An old reservoir, about 80 m. in diameter, 
presumably dating from when the abandoned 
cabins were occupied, receives most of the spring 
water, and a smaller pond was seen in the exten- 
sive grassy meadow below. A moist meadow 
extends to elevations at least 5 m. above the 
present level of the reservoir. The southernmost 
spring, 1 .6 km. by road south of the ranch houses, 
we found to be an open, bubbling spring hole of 
warm water ( 30 C\ ). Here and in open, shallow 
sloughs bullfrogs, but no fish, were seen. 

The tui chub, Gila hicolor. also occurs in the 
main spring, and is also represented by an endemic, 
somewhat dwarfed subspecies, G. h. isolata (pp. 
175-180). The tui chubs abounded, but the 
speckled dace, in contrast, appeared to be so 
scarce, and .so secretive, that it proved difficult to 
collect a good series. Not one specimen of Rhin- 
ichthvs was included in the second collection. The 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



135 



numbers of speckled dace are apparently being 
held down by introduced animals, especially large- 
mouth bass. Microptcriis salmoides ( 1 adult 
caught). An adult carp, Cyprinus carpio. was 
seen, and bullfrogs. Rami catesbeiona. were com- 
mon. In 1966 Stephen H. Berwiclc (class report. 
University of California, Berkeley, 1966) found 
Micropteiiis salmoides present, was told by a 
ranch worker that carp occur, was informed by 
William Nisbet, a state fishery biologist, that 
exotic fish were probably not present in I960, 
and that the bullfrogs, which abounded in 1966, 
were stocked by the state Department of Fish and 
Game about 10 years previously. 

Despite the abundance of water ( the main out- 
flow was estimated by us to be about 200-300 
gallons per minute ) . the local form of dace seems 
to be in the endangered category. 

The circumstance that largemouth bass and 
carp occur in Warm Springs led us to wonder 
whether the dace and the chub might have been 
introduced. However, the state authorities, when 
checked for us by Donald E. Lewis, then Manager 
of the Ruby Lake National Wildlife Refuge, could 
provide no evidence of such stocking. Later, 
Thomas J. Trelease, Chief of Fisheries in the 
Nevada Fish and Game Department, indicated to 
us that he could find no record that the minnows 
had been introduced. Distinctive characters also 
lead to the conclusion that both species are native. 

Description and Comparisons. 

Holotype, UMMZ 186905, an adult female 
35.3 mm. in standard length (fig. 24D). Para- 
types, UMMZ 186519, all other known speci- 
mens ( 100, 18-39 mm. long), including the adult 
male, 34. 1 mm. long, that is illustrated ( fig. 24C ) . 
All specimens are from the same Location, R12 
(data given above). 

Next to R. o. reliqmis (pp. 121-128), this is 
the most distinctive of the various local forms that 
appear from physiographic and systematic evi- 
dence to have been derived from Rhinichthys 
osculiis robtistiis. or from its immediate ancestor. 

In the following description, R. o. letlwporus 



is compared primarily with R. o. oligoponts, 
which occurs in the same pluvial lake basin. 

Size. This appears to be the most dwarfed 
among all the local forms of Rhinichthys from 
springs in the area under consideration. The 
largest specimens among the 101 collected mea- 
sure 34 mm. for males and 39 mm. for females, 
in standard length. The adults are not much 
larger than young of the year of some of the other 
forms. In degree of dwarfing, this form is closely 
approached only by the four warm-spring popula- 
tions of the Lake Diamond drainage basin referred 
to R. o. rohiistiis (table 20). 

Coloration ( fig. 24C.D ) . The general colora- 
tion is rather similar to that of R. o. oUgoporus. 
The primary band is rather sharp, at least pos- 
teriorly, and a considerable trace of the lower 
lateral band is evident on most of the specimens. 
The dark speckling is usually very fine and tends 
to extend downward across the caudal peduncle 
(in this respect contrasting with R. o. oUgoponis 
from Location Rll, but not .so strongly with 
those from RIO). Many of the specimens show 
a blackish wedge or streak along the lower edge 
of the caudal peduncle. In contrast with R. o. 
oUgoporus. at least a trace of a dark horizontal 
stripe, restricted largely to snout and upper part 
of opercle, is usually developed on the head, much 
as in R. o. rohiistiis. In the blackening of the 
crotches of the bifurcating branches of the rays 
in the dorsal and caudal fins, and occasionally 
the anal fin. this subspecies resembles R. o. robus- 
tiis more than R. o. oUgoporus. Only a few of the 
preserved specimens show a light streak above 
the main lateral band ( frequently shown by R. o. 
oUgoporus). In general, the color pattern agrees 
with that of the other forms. The black wedge on 
the middle of the base of the caudal fin is oc- 
casionally conspicuous, as it usually is in R. o. 
rohustus. but is commonly obsolescent, as in R. o. 
oUgoporus. In the fine speckling of the lower 
sides, as well as in the form of the body, this form 
of Rhinichthys somewhat simulates the young of 
Gila bicolor. 

Life colors. No bright colors were observed 
in the field. 



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CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Form (fig. 24C,D). The t'onn of this sub- 
species, unusually compressed for a Rhinichthys. 
is particularly distinctive: the greatest width of 
the body steps about 2.0 times into the depth over 
the curve of the sides, rather than about 1 .5 times 
as in R. <>. rohii.stiis (in this respect R. o. oligo- 
IHnns is approximately interniediale ). The an- 
terior profile is less flattened than in R. o. rohitstiis 
and is less arched than in R. o. oliiioponis. The 
anterior part of the head is more foreshortened 
than in R. o. robtistiis. but is rather more pointed 
(less rounded) than in R. o. olii^oponis. The 
mouth is definitely straighter than in R. o. 
oUgoporus and more oblique, rising forward to a 
horizontal through the lower edge of the eye. 

Fri:num. Not one of the specimens was ob- 
served to have a frenum. 

Barbisl (table II). This subspecies is one of 
the three under consideration that almost invari- 
ably lacks the barbel. Among 79 specimens only 
4 had a barbel on one side (minute in 3, small 
in 1 ), and none had a barbel on both sides. 

Lateral-line sYSTiiM (table 12; figs. 25. 26). 
This form again, as are R. o. rcliqitiis and R. o. 
oligoponis, is characterized by a great reduction 
in the development of the lateral line on the 
body, even more than in R. o. oligoponis. less 
extreme than in R. o. icUqmis. An average pore 
count of 1.0 is not attained until the standard 
length reaches about 32 mm. and only 1 count 
higher than 6 was obtained among 136 sides 
enumerated. 

The lateral-line system is also degenerate on 
the head. All 21 specimens examined for this 
character had the supratemporal canal inter- 
rupted. In this respect, the agreement is best with 
R. (>. rcUqiius and contrasts somewhat with R. o. 
oligoponts. In some specimens even the lateral 
pores of this canal are represented merely by 
neuromasts. 

Scale structure. The scales show the typical 
structure of Rhinichthys (p. 97 ). 

Morphometry (table 13 ). In correlation with 
its compressed form, as noted above, the body 



proper, and more strikingly the caudal peduncle, 
are on the average deeper in each sex than in the 
other forms here treated. The relatively long 
head and the particularly large eye reflect the 
dwarfing of this form. The other head measure- 
ments are not particularly distinctive. As usual in 
R. o. rohitslii.s and its derivatives, the fins are 
small and rounded. On the average, the dorsal 
and anal fins are inserted farther back than in R. 
o. robiistus, except for some of the populations 
isolated in the Diamond Valley drainage system; 
as far back, or farther, than in R. o. ohgoponis. 
As determined from radiographs, the pelvic in- 
sertion lies under the 15th vertebra, occasionally 
under the 14th or 1 6th. less far back than in R. o. 
rcliqiius ( table 17). 

Particularly striking is the strong obliquity of 
the nearly straight mouth, so that the upper jaw 
rises to about the level of the middle of the eye. 
In R. o. ohgoponis the mouth is more curved, 
and rises only to opposite the lower part of the 
eye. The difference in the angulation of the 
mouth is less striking in the young, which in all 
the subspecies of Rhinichlhys osctihis. and in 
many cyprinoid fishes, tend to have an oblique 
mouth, in correlation, presumably, with particu- 
late midwater feeding on small invertebrates — a 
frequent developmental phenomenon, even of 
the white sucker. Calo.stonuis coninicisonnii 
(Lacepede). as shown by Stewart ( 1926). This 
feature of R. <>. Iclhoporus, therefore, appears to 
be a trophic adaptation in which the juvenile 
character is retained by the adult. In the bottom- 
land Warm Springs of Independence Valley, 
where there has been considerable depth of rela- 
tively quiet, vegetated water, there would have 
been some advantage to midwater feeding through- 
out life, whereas in the shallow current on the 
alluvial slopes of the Clover Valley springs, in- 
habited by R. (>. oligoponis. bottom feeding would 
have been more in order. 

Sexual i^imorphism and nuptial charac- 
ters. Sexual dimorphism in the size of the fins 
(table 14) is relatively slight for the caudal and 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



137 



Table 15. Number of fin rays in populations of Rhinichthys osciilus in certain hasins in Nevada. 



Subspecies 

Pluvial lake system 
Locality 



Dorsal rays' 



9 Mean 



Anal rays' 



8 Mean 



Rhinicliihys oscnius rohiisiiis 
L. Lahontan (Humboldt R.) 

Carico L. Valley 

Crescent Valley 

Bishop Creek 

Springs near Town Cr. 
L. Franklin (introduced) 

Ruby Valley 
L. Diamond 

Potts Ranch 

Dianas Punch Bowl 

Coils Creek 

Birch Ranch 

Big Shipley Spr. 
Rliiniclitliys osciiliis rclicjinis 
L. Gilbert 

Grass Valley 
Rliinicluliys osciiliis oligoponis 
L. Clover 

9.5 mi. S. of Wells 

Warm Sprs., Clover V. 
Rhinichtliys osciiliis Iclhoponis 
L. Clover 

Independence Valley 



— 


3 


— 


8.0? 


4 


30 


— 


7.88 





74 


— 


7.97 


6 


98 


'> 


7.96 



13 



7.93 



_ 


52 


— 


8.00 


4 


90 


6 


8.02 


9 


31 


— 


7.77 


5 


30 


— 


7.86 


8 


28 


— 


7.78 


T 


77 


10 


8.09 


I 


31 


1 


7.77 


3 


38 


— 


7.93 



- 


3 


- 7.0? 


— 


34 


— 7.00 


1 


74 


1 7.00 


5 


100 


- 6.93 



13 — 



74 



2 40 
1 39 



6.93 



1 


56 


— 


6.98 


T 


92 


6 


7.04 


1 


38 


1 


7.01) 


1 


34 


— 


6.97 


4 


30 


1 


6.86 



1 7.94 



9 7.05 



1 6.98 
1 7.00 



6.95 



Subspecies 

Pluvial lake system 
Locality 



17 



Caudal rays" 



19 



20 



Mean 



Pelvic rays" 



9 Mean 



Rliiniclitliys osciiliis robustus 
L. Lahontan (Humboldt R.) 

Carico L. Valley 

Crescent Valley 

Bishop Creek 

Springs near Town Cr. 
L. Franklin (introduced) 

Ruby Valiey 
L. Diamond 

Potts Ranch 

Dianas Punch Bowl 

Coils Creek 

Birch Ranch 

Big Shipley Spr. 
Rhiiiichlhys osculiis reliqiiiis 
L. Gilbert 

Grass Valley 
Rliiniclitliys osciiliis oligoponis 
L. Clover 

9.5 mi. S. of Wells 

Warm Sprs., Clover V. 
Rliiniclitliys osciiliis letlioporiis 
L. Clover 

Independence Valley 



- 


1 


— 


1 20.0? 


5 


27 


— 


— 18.84 


3 


32 


-) 


— 18.97 


S 


-n 


1 


I 18.70' 



1 — 



13 — 



18.93 



— 


30 


— 


— 


19.00 


11 


85 


I 


— 


18.83 


5 


34 


1 


— 


18.90 


— 


27 


-> 


— 


19.00 


1 


29 


-> 


— 


1 9.00 


5 


54 


9 


— 


19.00 


3 


36 


-> 


1 


19 07 


3 


33 


4 


1 


19.07 


3 


28 


1 




18.94 



1 — 



1 


5 


— 


7.8? 


7 


59 


'■} 


7.97 


18 


130 


— 


7.88 


67 


145 


— 


7.68 



141 



7 64 

6 86 



103 



36 



7.59 



39 


83 


— 


7.68 


72 


124 


9 


7.63 


71 


281 


~t 


7.79 


129 


37 


— 


7.22 


123 


39 


— 


7.23 



— 7.18 



6.95 
6.96 



6.97 



1 Last 2 elements counted as one ray. 
- Principal rays: branched rays -+- 2. 
^ Both sides counted in most specimens. 

* One count of 14 in an apparently normal, uninjured fin; included in calculation of mean. 
^ One count of 16 in an apparently normal, uninjured fin; included in calculation of mean. 

(Tabic contmued on next page. ) 



138 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 15. continued. 



Subspecies 

Pluvial lake system 
Locality 



Pectoral rays' 



10 



11 



12 



13 



14 



15 



16 Mean 



RItinichthys osciiliis rohiisliis 
L. Lahontan (Humboldt R.) 

Carico L. Valley 

Crescent Valley 

Bishop Creek 

Springs near Town Cr. 
L. Franklin (introduced) 

Ruby Valley 
L. Diamond 

Potts Ranch 

Dianas Punch Bowl 

Coils Creek 

Birch Ranch 

Big Shipley Spr. 
Rhiiiichlliys ti.sciiliis rclicjims 
L. Gilbert 

Grass Valley 
Rliiiiiciilhys ti.wiiliis i)lit;i_)ponis 
L. Clover 

9.5 mi. S. of Wells 

Warm Sprs., Clover V. 
Rhiiiichlliys osciiliis tclhoporiis 
L. Clover 

Independence Valley 



— — 6 



— 4 



4 


T 


— 


— 


— 12,3? 


3 


21 


28 


16 


— 13.84 


3 


20 


33 


14 


— 13.83 


7 


31 


52 


8 


1 13.65 



53 



16 
19 



16 



10 



35 



12 



65 15 — 



31 



I 13.89 



7 


45 


31 


3 


— 


13.35 


9 


82 


78 


29 


■> 


13.67 


7 


24 


40 


9 


— 


13.64 


13 


30 


19 


8 


— 


13.31 


11 


31 


23 


4 


— 


13.29 



12.65 



— 12.49 

— 12.72 



12.72 



Table Ifi. Number of vertebrae in pt>piihnioiis of K\\mKh\hys oscu\u^ in cerlain basins in Nevaclu.^ 



Subspecies 

Pluvial lake system 
Locality 



Precaudal 



Caudal" 



TotaP 



18 19 20 21 22 Mean 



16 17 18 19 20 Mean 



35 36 37 38 39 40 Mean 



R 



1. 



robiistus 

Lahontan (Humboldt R.) 

Carico L. Valley 

Crescent Valley 

Bishop Creek 

Springs near Town Cr. 

f-ranklin (introduced) 

Ruby Valley 

Diamond 

Potts Ranch 

Dianas Punch Bowl 

Coils Creek 

Birch Ranch 

Big Shipley Spr. 
reliqiuis 
L. Gilbert 

Grass Valley 
R. o. oligopnrns 
L. Clover 

9.5 mi. S. of Wells 

Warm Sprs.. Clover V. 
R. i>. lelhoponis 
L. Clover 

Independence Valley 



R 



- 


— 21— 20.3? 


-> 


18.0? 


1 


1 — 38.5? 


— 


9 18 2 — 19.76 


— 2 21 


6 — 18.14 


6 20 


3 — 37.90 


5 


12 18.71 


— 2 12 


3 — 18.06 


— 6 11 1 


36.72 


_ 


12 7 19.35 


— 1 8 


4 — 18.23 


7 5 


1 — 37.54 



19.64 



18.55 



10 7 — 


— 19.26 


1 12 4 


17.18 


8 6 — 


— 19.00 


— 2 11 


5 2 18.35 


4 13 2 


— l^t.85 


— 7 9 


17.44 


18 11 — 


1 19.29 


— 14 15 


2 — 17.61 


10 10 1 


— 19.56 


— 5 13 


1 — 17.84 



— 1 5 3 — 38.22 

7 10 36.59 

2 10 4 4 — 37.50 

— 961— 37.50 

8 17 5 — I 37.00 
1 9 4 37.42 



II 16 10 — 19.97 



— 2 18 12 — 20.31 

— 4 12 5 — 20.05 



I 9 10 



19.45 



21 13 3 16.51 



12 18 2 16.69 

2 10 5 2 — 17.37 



111 5 17.24 



3 16 15 3 



36.49 



4 24 4 37.00 

3 6 8 3 — 37.55 



7 11 1 



36.68 



^ All counts from radiographs. 

- Including hypural complex (as 1) and the 4 vertebrae comprising the Webenan apparatus. 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



139 



Table 17. Position of pelvic-fin insertion, in terms of overlyinc; vertehrii. in populations of Rhinichthys osculus 
in certain basins in Nevada. 



Subspecies 

Pluvial lake system 
Locality 



13 



14 



Vertebra no. (from radiographs) 



15 



16 



17 



18 



Mean 



Rliinichthvs osculus rohiistiis 










L. Lahontan (Humboldt R. ) 










Carico L. Valley 


— 


— 


1 


2 


Crescent Valley 


— 


1 


17 


14 


Bishop Creek 


1 


3 


12 


1 


Springs near Town Cr. 


4 


9 


6 


— 


L. Franklin (introduced) 










Ruby Valley 


— 


*> 


6 


3 


L. Diamond 










Potts Ranch 


1 


7 


10 


-> 


Dianas Punch Bowl 


— 


6 


7 


7 


Coils Creek 


— 


— 


12 


8 


Birch Ranch 


2 


8 


11 


— 


Big Shipley Spr. 


1 


3 


10 


6 


RhinichlhYS osculus reliqiiiis 










L. Gilbert 










Grass Valley 


— 


— . 


2 


21 


Rhiniclilhys osculus oligoporus 










L. Clover 










9.5 mi. S. of Wells 


— 


1 


11 


7 


Warm Sprs., Clover V. 


— 


2 


12 


11 


Rhiniclitliys osculus letlioporus 










L. Clover 










Independence Valley 


— 


3 


16 


3 



15 



15.7? 
15.45 
14.76 
14.11 

15.09 

14.65 
15.05 
15.40 
14.43 
15.05 



16.38 



15.32 
15.36 



15.00 



Table 18. Nuiuher of scale rows and gill-rakers in populations of Rhinichthys osculus in certain basins in Nevada.^ 







Scale 


rows 






Subspecies 






















Pluvial lake system 


Lateral-line 


Predorsal 


Around 


Around 


Gill-rakers 


Locality 


series 


series 


body 


peduncle 


(total) 


R. o. robuslus 












L. Lahontan (Humboldt R. ) 












Carico L. Valley 


69-78(73.7;,) 


38-43(40.0:,) 


59-64(61.7:,) 


30-34(31.7.,) 


6-8 (7.0?:,) 


Crescent Valley 


66-75(70.5-.,,) 


40-47(42.9=,,) 


52-72(66.8:,,) 


32-40(35.9=„) 


6-10(7.30-.,,) 


Bishop Creek 


59-70(63.8.«,) 


35-41 (38.3is) 


57-68(61.8,,,) 


30-33(31.3,:) 


5-9 (7.30:,,) 


Springs near Town Cr. 


62-70(66.8.v) 


33-46(40.7„) 


58-70(64.7=,,) 


28-34(32.2,,,) 


6-9 (7.35-3,) 


L. Franklin (introduced) 












Ruby Valley 


61-69(64.9,,) 


33-43(38.9,,) 


60-66(63.7,,) 


32-36(33.4,,) 


6-9 (7.47,,-.) 


L. Diamond 












Potts Ranch 


54-64(59.3:,,) 


33-41(35.8..,,) 


56-67(61.4,,) 


30-34(32.1,,) 


6-9 (7.41=) 


Dianas Punch Bowl 


56-72(61.8,,,) 


31-44(36.8,:,) 


54-69(59.3,:,) 


29-33(30.6,.,) 


6-9 (7.50.,,) 


Coils Creek 


58-72(65.5,,,) 


37-44(40.3,,,) 


60-70(64.4,,) 


31-34(32.3,,,) 


5-10(8.05,b) 


Birch Ranch 


52-65(57.9,.:,) 


32-40(34.7=3) 


5.'i-62(58.7..„) 


28-32(30.7-..,) 


6-10(7.33=:) 


Big Shipley Spr. 


52-71(58.7,,,) 


32-38(34.82,,) 


53-60(55.7,:,) 


28-32(29.7,,,) 


7-10(7.65:,,) 


R. o. reliquits 












L. Gilbert 












Grass Valley 


60-74(66.4:.,) 


37-47(41.6:.:) 


56-76(66.5=) 


32-40(36.2-..,) 


6-9 (7,21:«) 


R. o. oligoporus 












L. Clover 












9.5 mi. S. of Wells 


54-60(57.5.) 


35-38(36.7,,) 


58-62(59.8:,) 


29-32(30.2.,) 


7-9 (8.00,) 


Warm Sprs.. Clover V. 


54-66(59.3,.) 


34-40(37.3,5) 


52-60(55.5,:,) 


28-31(29.9,,,) 


5-9 (7.07,,-,) 


R. o. letlioporus 












L. Clover 












Independence Valley 


50-62(56.3..,) 


26-35(29.8=,) 


48-60(53.5,n) 


26-34(29.7,,,) 


6-10(7.90=,,) 



^ For each entry there is given the observed range and, in parentheses, the mean, with the number of specimens as a subscript. 



140 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 19. Sex ralias by number and by /nonuiss for 
popuhilicns of Rhinichthys osculus //; certain basins in 
Nevada.'^ 



Subspecies 

Pluvial lake system 
Locality 


Sex 


ratio by 


M per 
100 F 


Biomass, 

M/F 


R. <i. roliiisiiis 






L. Lahontan ( Humlmklt R.) 
Crescent Valley 
Bishop Creek 
Springs near Town Cr. 


47 
67 
54 


.30 
.48 
.38 


L. Franklin (introduced) 
Ruhy Valley 


75 


.26 


L. Diamond 
Potts Ranch 
Dianas Punch Bowl 
Coils Creek 
Birch Ranch 
Big Shipley Spr. 


124 
80 
52 
96 

25 


1.10 
.69 
.36 
.63 
.12 


R. o. rcliqiins 






Lake Gilbert 
Grass Valley 


63 


.27 


R. i>. <ilii;oi^oriis 






Lake Clover 

9,5 mi. S. of Wells 

Warm Sprs.. Clover Valley 


75? 
93 


.6.S 
.40 


R. (>. Iclluiponis 






Lake Clover 
Independence V. 


80 


.65 



' Bosed on detailed dula in table 20. 

the pelvic, alxnil nieditini for the diirsal. among 
the highest for the pectoral ( averaging 5.2 percent 
of the standard length longer in males than in fe- 
males). This high dimorphism in the size of the 
pectoral fin indicates that we are dealing with 
dwarfed adults, not merely with young specimens, 
and confirms that we are describing a distinct 
form. The single August 25 collection has not 
provided specimens in nuptial condition. The 
sexes differed little in size ( see below ) . 

Mi-RisTKs (tables 15. 16, 18). Dorsal rays 
usually S. occasionally 7, very rarely 9; anal 7, 
occasionally 6. The pectoral rays, as in subspecies 
R. o. reliijiius and R. o. (>lii;i>p(>nis. average few 
( 12.72 for 5(S counts). The pelvic-ray counts, 7, 
with variants of 6 seemingly exceeding those of 8. 
also yield a low average. The principal caudal 
rays are typically 19, but vary from 1 8 to 20. The 



vertebral counts average 19.45 precaudal. 17.24 
caudal, and 36.68 total, slightly fewer than in R. 
(>. oUgoponis. The scale counts, averaging 56.3 
in lateral-line series. 29.8 in predorsal rows, 53.5 
around body, and 29.7 around peduncle, are usu- 
ally fewer than in any of the other forms under 
treatment ( table 18). The trend toward low ray 
counts seems to be related to the dwarfing of this 
form. The gill-raker counts, however, average 
slightly on the high side ( table 18). 

PiiARYNCiiiAL TLi'TH. The 5 specimens counted 
have the expected tooth formula, I, 4 — f, 1. The 
single tooth of the lesser row is strong. 

Se.XUAL i:)IIFERENCES IN NUMBERS AND BIO- 
MASS (tables 19, 20). This form is greatly 
dwarfed, to the extent that it has been apparently 
impossible to .separate young from adults by size. 
Collecting in the very dense vegetation was so 
difficult that some young probably escaped, al- 
though the fish were taken by a fine-meshed 
woven ( "Ct)mmon Sense") seine, as well as by a 
tied-mesh seine, and special effort was expended 
to secure a representative sample. The compressed 
grouping, with males predominating in the lower 
size groups, suggests that at most only a few of 
the larger young were seined. All specimens were 
sexed by gonad examination. The 101 specimens 
that we were able to collect comprised 45 males 
18-34 mm. in standard length, mostly in the 
2.5-imii. and 3.0-mm. size groups, and 56 females 
21-39 mm. long, mostly in the 2.5-mm. to 3.5- 
mm. size groups. The average standard lengths 
of the sexes were extremely close, 26.7 and 30.1 
mm., for the males and females, respectively. The 
sex ratio, 80 males to 100 females, is much closer 
tt) eqtiality than in the iilhcr poptilations, as is also 
the ratio of males to females by bulk, .65: 1 . The 
close similarities between the sexes in numbers, 
mass, and size presumably reflect the extreme 
dwarfing, and suggest that this subspecies may be 
an amuial fish. 

Derivation of name. The name Icihoponis 
was derived, figuratively, from Aj/^//, a forgetting, 
and 7r(7».s, a passage through the skin. It is re- 
garded as in adjeclival form. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



141 



Dh 


q 


V3 


nf 


<4-l 











W^ 


Ui 




<u 


Ct 


Xi 




B 


O 






J e 

t/iE 



t/5 



O . 
^1 



a 3 ■ 
« — I 

JDO. 



ON n-i ri — 
— — f^ 'T 


1 -t r~~ 


1 ON OS O ^O 
1 ^ rJ r-- 


ON r~ 


rr, -t r*-, 0\ 

ri — r<-, ri 


|C,- 


1 ri O ON r*-. 
1 r'. n r^. T-~ 




— CO ^ OS 


^ 1 1 


1 00 W-. r^. r*-. 
1 ri n-- ri r^. 


-T ri 



Tj- ri m — r- 



- I 



^D'^oq-^-^v^-tr-"^ 


•^ CO O 


r]0\*^^ 


O O CO o" O '^' O ^' ND r-" 


O 00 Tf' 


r- m Tj- 

r-, r*i V, 



I I I I 1 I I 1 I III 

r^ r*-, rJ >/"* ON NO Vt — O -T" r^, r^-i 

«/-i — r*-, r^i ri rj ^^ r'-. r*-. — ' C I ri 



Of O Ot Ot 

r^. ONON"^-^— ^OfNr*-, Ov ■^^^^£) 



-1- o 



m "^ "^ -Tf r- 



" "1 CO "^ I tn 00 '^ 00 
(N NO r*-i 



f*", -T r*-i -r r I »/-i r^ r I -i- no r<-. vi 

I I 1 I I I I I I I I I 

ONTiri — f. riTj-r-rjOmri 

— ri ri ri — fo fo — ri ri r^j r) 



Of Of 

*00f'^0+*^^0f'O*O0t'T30t 

r-vo^r-^os — NONOON<^rim 

*r^ -rf OO <0 f~^ 0\ ^C OCONOOm 



r- NO NO o T- -t 

On r*~i r^, fi ri r-- 

ri — ^ ^ r-, 



c — in 00 "^ ON 

r^. -^ v^ n -+ >/-. 

I I I I I I 

"^ O — ■ n oo oo 

— -7f >n — 1 rJ ri 



Of Of 

•so -SD Ot *<) *0 Of 

> -^ o rj NO 



I 1 

00 — 

— rt 



> G 

> 



— lo ^H r*^ 



^- '3N « 



f^ 



oo 



CT\ — — 




OS 

2 
o 

- = 

5 1 ° 
: o u 



c 

o 
H 



o 

03 



O 



o. 

C/2 






O ™ a 
-JO 



U 

O 



s > 



o 
0. 



o 



o 



Oi 









">■ 


o 






— 






— 
a, 


^ 


> 





c/i 


IS 


^ flj 




<i> 




1/5 


: 




; 


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ON 



o > 
"5 .2 
.2 U 



142 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



TUI CHUB 

Gila bicoloi (Girard). 

Under this species name we treat all native 
chubs that occur in the area of Nevada under 
report (pp. 1-2), along with the two com- 
mon forms, 'pcctiuijer and 'obcsa.' of the central 
part of the Lake Lahontan system. Different types 
from disjunct basins are referred to several distinct 
subspecies ( table 21). in view of the complicated 
interrelationships of these forms, and their con- 
fused classification and nomenclature, some con- 
sideration needs first to be given to the Lahontan 
forms. Although we have long had other related 
forms under study, we refrain in this report from 
treating the several unnamed subspecies that we 
have indicated ( Hubbs and Miller, 1948b, p. 91) 
as occurring in different springs in the drainage 
basin of pluvial Lake Railroad, which lies within 
the area on which we are reporting, as well as 
various extralimital subspecies that occur in other 
endorheic basins in the drainages of pluvial lakes 
Toiyabe ( </. v.). Dixie, and White Mountains in 
western Nevada, as well as various forms that 
occupy, or until recently occupied, lake basins in 
California, southeastern Oregon, and southeast- 
ern Washington (Hubbs and Miller, 1948b, pp. 
43_45, 61-67. 7(_)-74; Miller, 1973. pp. 1-8). 

The populations of Railroad Valley have been 
referred by us (Hubbs and Miller, 1948b, p. 91 ) 
to Sipluilcles ohesiis and by Bradley and Deacon 
(1965. p. App. II- 1 ) to Gila hicalar. A wide- 
ranging subspecies t)ccurs in Little Fish Lake, in 



Twin Springs near the outlet end of Hot Springs 
Valley, and in the Duckwater Creek tributary to 
Railroad Valley (also in Artesian Well No. 7 in 
the playa of Railroad Valley, presumably as a 
result of floodwatcr incursion from either Twin 
Springs or Duckwater Creek ) . That form we have 
found to occur also, presumably as a result of 
stocking from Twin Springs, in Stone Cabin (or 
Willow Creek) Valley, just to the southwest, 
across a definite divide (p. 232). 

Gf.ni^ral Appraisal Of Lahontan Subspecies 

Several complications arise in the classification 
and nomenclature of the forms listed in table 21. 
The generic pertinence of the species calls for 
discussion first. Until very recently, the species 
complex involved has been treated under the 
generic name Sipliaiclcs. On the basis of Teruya 
Uyeno's doctoral study (1960), Siphatcles has 
been synonymized with Gila. Preliminary notices 
to that effect have been given by Miller ( 1961, 
p. 384; 1968, p. 171 ) and by Bailey and Uyeno 
( 1964). and the species name 'hicolor is listed 
under Gila by Bailey el al. ( 1960, p. 13; 1970. 
p. 20). The basis for this action has been ( 1 ) 
the indication of the very close agreement be- 
tween "Siphatcles obcsiis" {Gila hicolor ohesa) 
and the Utah chub. Gila atraria (Girard) in ex- 
ternal and. especially, in skeletal characters, and 
( 2 ) the opinion that these points of agreement 
overbalance the difference in dentition that pre- 
viously had been accorded greater weight. The 



Table 21. Local fanus af Gilii hicolor williiii ihc area <</ //;<■ picscnl study ami in llic central part <</ the Lahontan 
system} 



Subspecies 



Pluvial 
lake system 


Valley 


Lahontan 


Humboldt R., etc. 


Lahontan 


Humboldt R., etc. 


Dianionil 


Diamond 


Newark 


Newark 


Newark 


Fish Creek (Little Smoky) 


Clover 
Railroad 


Independence 

Railroad; Warm Springs; Little Fish Lake 



Gila hicolor pcclinifer (Snyder) 

Gila hicolor ohcsa (Girard) 

Gila hicolor ohaa (2 aberrant stocks) 

Gila hicolor /ic vi«rAi'».s7v 

Gila hicolor euchila 

Gila hicolor isolata 

Gila hicolor: several unnamed subspecies, still under study 



•The four subspecies other than C. b. peclinijer and those of the basin of Lake Railroad are listed in more detail on pasie 15" and are treated 
in detail in this report. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



143 



circumstance that the pharyngeal teeth of Sip- 
luiteles are invariably uniserial, whereas in all 
species of Gila (sensii stricto) these teeth are 
biserial, was discounted, because the same differ- 
ence frequently exists within other recognized 
genera and even within species, and the difference 
in dentition may have merely trophic implications. 
However, the whole problem of generic limitations 
in the Holarctic Cyprinidae is presently in almost 
utter confusion. We merely adopt for the present 
discussion what seems to be the current trend. The 
decision has little effect on the problems of the 
present report, except for the indication that the 
Lahontan and Bonneville species, respectively 
^bicolor and 'atrarUi,' are now believed to be 
much more intimately related than was signified 
by their generic separation. 

Some of the osteological features of Gila bicolor 
are briefly indicated in the diagnosis of the genus 
Relicltis (pp. 182-193). 

Species name. The specific name bicolor is 
accepted in place of obesa on the basis of a 
nomenclatural muddle that Bailey and Uyeno 
( 1 964 ) clarified. The name obesa remains avail- 
able, and has been accepted, for a Lahontan sub- 
species. 

Status of the lacustrine type (pcctinijci). 
Problems of taxonomy as well as nomenclature 
are involved in the treatment of 'pectinijcr as well 
as 'obesa' as subspecies of Gila bicolor. In de- 
scribing Leucidius pectinijer, Snyder ( 1917, pp. 
60-67, figs, 4-6) di.stinguished this form as a 
distinct genus on the basis of its very numerous 
gill-rakers, 29 to 36 and its tooth formula, 5 — 5 
vs. 5 — 4. However, as we note in the discussion 
(pp, 146-149) of the gill-raker character, there 
is a complete gradation between topotypic 'obesa,' 
with 1 1 to 19 rakers, and 'pectinijer,' with counts 
to 40, Furthermore, isolated populations show all 
levels of raker number, as we have shown in 
studies hopefully leading toward a revision of the 
whole group. Some specimens of 'pectinijer' 
have 5 — 4 teeth and some of 'obesa' have 5 — 5 
teeth. The proportion of specimens of the 'obesa' 
type with 3 — 5 teeth is highest in series, like that 



from Carson River near Fallon and from the 
Humboldt River near Lovelock (table 23), that 
apparently show introgression from 'pectinijer 
also in the increased and skewed raker number 
(table 22). 

The striking increase in raker number and 
length in 'pectinijer' is paralleled again and again 
among cyprinids and other freshwater fishes, 
throughout the world, as they have assumed a 
lacustrine existence. The structural modification 
is clearly related to the well authenticated notable 
abundance of plankton in lakes and general 
paucity in streams. The less robust teeth of 
'pectinijer,' numbering 5 — 5, probably constitute 
a parallel adaptation. 

Essentially, 'pectinijer' and 'obesa' appear to 
be trophic adaptations with, in general, ecological 
segregation (which has been decreasing, with the 
agriculture-based desiccation of the waters). As 
indicated in the discussion of the gill-raker char- 
acter (pp. 146-149), the high number and 
greater length of the rakers in 'pectinijer' is inter- 
pretable as an adaptation to plankton feeding in 
the lakes it inhabits, currently the remnant lakes 
of the Lahontan basin. We may assume that it 
abounded throughout Lake Lahontan, as Snyder 
( 1 9 1 7. p. 66 ) found it to swarm in Pyramid Lake, 
"approaching the shore at times in enormous 
schools," that "resemble large purple clouds." In 
Lake Lahontan it was presumably almost com- 
pletely segregated from the fluviatile form 
( obesa ) . 

After noting that Leucidius pectinijer had 
"been reduced to a subspecies of S. bicolor, of 
which it is considered to be the lacustrine form," 
La Rivers and Trelease ( 1952, p. 117) put forth 
the following proposition: 

However, on the basis of recent data, we are 
convinced that pectinijer has no valid standing 
as a taxonomic unit. Genetically, pectinijer 
might be preserved to indicate tui chub with 
fine gill rakers (as Shapovalov and Dill. 1950, 
have done), just as it might be feasible, under 
some circumstances, to so distinguish between 
people with blue eyes and people with brown 
eyes. Chub with coarse gill rakers occur side-by- 



144 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



side with individuals with line gill rakers, con- 
trary to Snyder's supposition that he could oh- 
serve them segregating in separate schools in the 
lake. Gill net sampling in Pyramid Lake hy 
the writers has resulted in catches of chuhs 
with hoth types of gill rakers at the same time 
and place, and there is no sexual correlation 
between raker differences. Whether there are 
any differences in feeding habits between the 
two types remains to be seen, but the above 
sampling, dining the winter, showed both to 
be mixed in the same schools and to be feeding 
on the bottom on the same materials. It is 
possible that at other times of the year one 
form may have an advantage over the other in 
being able to obtain more plankton from the 
water, but at present, that i^ a dubious point. 

Later ( 1962, pp. 412-421, figs. 192. 193) La 
Rivers expanded on the same line, obviously not 
cognizant of the large stream populations of G. 
hicolor obcsii. He still felt "convinced that pectini- 
jcr has no valid standing as a taxonomic unit." 

We regard as wholly untenable the view that 
'ohc.siis" and "pcctiuijcr should not be distin- 
guished. That view was not supported by Shapo- 
valov and Dill. who. on our recommendation, 
merely listed the two forms as subspecies of the 
tui chub, "Siplnitck's" hicolor. Snyder collected 
sizable .series (he listed Kl counts of scales). He 
took ^pccliiiilcr' only in lakes and he did give 
some evidence ( p. 67 ) of segregation. The two 
types differ not only in the number and length of 
the rakers, but also in dentition, body and head 
form, and usually in scale number. They do. 
however, intergrade, and probably have done so 
increasingly as surface waters have diminished 
through agricultural use. 

For about three decades LciicicUiis has been 
treated as a synonym of Sipluilclcs and by some 
authors S. pcctinifcr and S. ohcsiis have been inter- 
preted as only subspecifically distinct, primarily 
on the basis of apparent trophic adaptation and of 
extensive introgression and intergradation. with 
occasional fusion ( Hubbs, 1941, p. ISS; Hubbs 
and Miller, 1943, p. 352; 1948b, pp. 41-42; 
Shapovalov and Dill. 1950, p. 386). The es- 
sentially undocumented interpretation was some- 
what elaborated later (Hubbs. 1961. pp. 13-14). 



Some further indication of introgression is fur- 
nished in the present report, and much further 
confirmatory data have been gathered by us in a 
survey of variation in Sipluiiclcs throughout its 
range. 

Because of the tendency toward sympatry of 
the two types in certain large lakes, particularly 
Lake Tahoe, Richard Ci. Miller (doctoral thesis. 
Stanford University. 1951) and Hopkirk and 
Behnke ( 1966. p. 136) have, in strong contrast 
with the opinion of La Rivers, interpreted \ibesa' 
and 'pc'ctinifer' as .specifically distinct. However, 
our studies have provided evidence of wholesale 
introgression and hybridization wherever the two 
types have met. In Eagle Lake, California, fusion 
of the two types has apparently led to the origin 
of a taxon with a distinctly bimodal, intermediate 
number of rakers ( Kimsey, 1954, pp. 397-398, 
fig. 2). though essentially uniform in the other 
taxonomic characters. 

We strongly suspect that where both occur in a 
lake the two types are segregated by schools and 
by habitat. Snyder, as noted above, wrote of huge 
schools of 'pcctinifcr' in Pyramid Lake, and we 
have studied in detail (Hubbs, 1961, pp. 13-14) 
a large collection from the inlet end of Walker 
Lake that was uniformly typical of 'pcctinifcr in 
all characters, except that a considerable propor- 
tion had. as an apparent result of introgression, the 
low raker number of 'obc.\(i.' and others had a 
dribbling range of counts fully connecting the 
high 'pcctinifcr' and the low 'ohc.sa' modes. 

Admittedly, the interplay between 'pcctinifcr' 
and 'obcsa could be interpreted as extensive in- 
terspecific hybridization rather than as subspecific 
intergradation. It is one of many cases in which 
the distinction is to a large degree arbitrary, such 
as one of us (Hubbs, 1943) indicated long ago. 
We favor the view that the extensive interchange 
in genes is better interpreted as intraspecific than 
as interspecific. Our reasoning parallels that ad- 
vanced by us (Miller and Hubbs, 1969) for re- 
taining the commoner of the widespread forms of 
the threespine stickleback as subspecies of a 
single species. Gnslcrostcu.'i aculcntus Linnaeus. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELIC T FISHES 



145 



Table 22. Nimiher of gill-rakers in populations of Gila bicolor in certain basins in Nevada. 

Subspecies Gill-rakers, including all rudiments, on first gill-arch 

Pluvial lake system 

Locality' 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 No. 

Gila bicolor obesa 
Lake Lahontan 

Carson River (Gl)= _ _ _ _ 1 6 9 13 13 8 5 ? 5 13 1 70 

Humboldt River 

Near Lovelock (G2) _ _ _ _ 3 i6 32 37 29 15 5 2 2 2 2 — 145 

NearCarlin (G3) _ __ _ 2 5 23 16 13 — 1 __ _ ___ 60 

Bishop Creek (G4) — — 1 16 66 83 25 1 ________ _ 192 

Lake Diamond 

Birch Ranch (G5) — — — — 9 28 22 9 2 — — — — — — — 70 

Sulphur Spring (G6) ____ 5 22 43 12 — — — — — — — — 82 

Gila bicolor newarkensis 
Lake Newark 

Near Diamond Peak (G7) — 1 8 23 38 22 5 3 — — — — — — — — 100 

Moores Ranch (G8) — — 2 14 39 34 1 1 — — — — — — — — — 100 

Warm Springs (G9) ___ 2 6 — — — — — — — — — — — 8 

Gila bicolor eiichila 
Lake Newark 

Fish Creek Springs (GIO) — — 13 40 56 24 5 ________ _ L38 

Gila bicolor isolala 
Lake Clover 

Independence Valley (Gil) 2 9 30 53 36 18 !_________ 149 

^ Expressed as numbered Locations in the G series. 
- Some introgression from Gila bicolor pectinijer. 



Ave. 



16.43 

14.90 
13.62 
12.61 

13.53 
13.76 



11.99 
12.23 
11.75 



11.77 



11.14 



One example of apparent introgression from 
G. b. pectinijer into G. b. obesa is furnished by 
our analysis of a sample of subtypical \ibcsa from 
the lower reaches of Carson River, one of the 
areas in which we find the numbers of gill-rakers 
increased and markedly skewed in distribution 
(table 22). The same lot shows a somewhat in- 
creased proportion of specimens with the 5 — 5 
dentition characteristic of 'pectinijer' ( see below, 
and table 23 ) . Yet the very slight positive correla- 
tion between raker and tooth counts ( table 24 ) 
seems to indicate essential integration of the 
stock, following some degree of long-past hybrid- 
ization. In the apparently well integrated form of 
presumed hybrid origin, in Eagle Lake (men- 
tioned above), the proportionate number of speci- 
mens with 5 — 5 teeth is about the same in the 
modal groups with rather few and rather many 
rakers: 5 among the 28 with 12-19 and 5 among 
the 33 with 23-30 rakers (data from J. B. 
Kimsey's A. M. thesis. University of California, 
Berkeley); apparently there is some genetic 
segregation in the inheritance of gill-raker number 



in Gila bicolor. as evidenced also by the case 
mentioned by Hubbs ( 1961, pp. 13-14). 

The tooth and gill-raker counts throughout the 
Humboldt River system and in the basins disjunct 
therefrom provide almost no evidence of introgres- 
sion from G. b. pectinijer in this area, except in so 
far as the upstream downward dine in gill-raker 
number (table 22) may be construed as such 
evidence. It is plausible to infer that the upper 
Humboldt River, from which, presumably, the iso- 
lated basins of pluvial lakes Diamond, Newark, 
and Clover originally received their Gila bicolor 
stocks, was populated during some Pleistocene 
time by G. b. obesa rather than by G. b. pectinijer. 
The circumstance that lacustrine types with a high 
number of rakers apparently did not evolve in any 
of these three ancient lakes suggests that such 
adaptation is not particularly rapid and/or in- 
evitable. Of course, it is remotedly possible, 
though seemingly improbable, that rakers did in- 
crease in number in the ancient lakes, but reverted 
( through reverse adaptation ) to the low number 
after the fish had become limited to springs and 



146 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




FiGUKE 2S. First gill-arches, with gill-rakers. in three races (two subspecies) of Gila (Sipliatelcs) hkolor from 
separate hasins in Nevada, based on adults of comparable size. Scales represent 1.(1 mm. Drawn hy Martha B. Lackey. 
A. G. h. (iht'sa. considered topotypic: Humboldt River near Lovelock (Location G2); LIMMZ 124873, no. 5: 90 
mm. B. O'. />. (ihcsii. aberrant race: Sulphur Spring, Diamond Valley (G6); UMMZ 1 24')2X, no. 8; 88 mm. C. G. 
h. iicwiii kin\i\. paratvpe: spring in Nev\ark Valley near Diamond Peak (Ci7l: UMMZ 1.^21X.s, no. 8; 8.S mm. 



spring creeks. However, in the Mohave River sys- 
tem, a lacustrine type v\'ith many rakers, whieh we 
now treat as GiUi hicolor mohavciisis (Snyder), 
has persisted under postpluvial conditions that are 
conducive to fluviatile types (Hubhs and Miller, 
1 943 ). in the Lake Railroad system, similar types 
with moderately numerous rakers have persisted 
in i.solated springs. 

Number and lhngth of (;iiJ,-R4Kr;Rs. The 
degree of development of rakers determined by 
methods outlined on pages 92-93, and ex- 
pressed by their total number and by their con- 
cordant length (tables 22. 23), provides some 
of the most significant indications of differentia- 
tion among the isolated stocks of Gila (Siphatclcs) 
hicolor in the basins under consideration. This 
was to be expected, since gill-raker development 
is closely correlated with size and type of available 
food and often differs markedly in different 
stocks, yet is widely recognized as being usually 
fixed genetically. 



Within stocks, and more loosely between 
stocks, the number and the form of the rakers are 
correlated, so that the difference in appearance is 
accentuated. When very few, as in G. h. new- 
iirkciisis (fig. 2SC), the rakers are usually very 
thick, opaque and fleshy, nearly smooth, and 
nearly or quite in juxtaposition basally. whereas 
in topotypic G. b. ohcsa (fig. 28A ) the rakers 
are rather slender, translucent and bony, mod- 
erately denticulate or crenulate on the inner edge, 
and generally well separated at the base. This 
contrast is particularly sharp between G. b. 
iicwaikoisis and the Sulphur Spring race (fig. 
28B) of G. b. obc'sa. 

The number of rakers in Gila bicolor varies 
extraordinarily, from 8 to 40. without any hiatus, 
and with many modes represented among the 
multitude of local forms we have had under study, 
not only in the Humboldt River system and the 
once connected endorheic basins (table 1 ). but 
also generally in the isolated waters in central and 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



147 



northern Nevada, central and eastern California, 
southeastern Oregon, and southeastern Washing- 
ton. As explained above, some distinct and geo- 
graphically separated forms have similar counts. 
The contrast between the subspecies is accentu- 
ated, for it involves also the length and crowding 
of the rakers — as was shown by Snyder ( 1917, 
figs. 4-5) in contrasting the rakers of the forms 
he called Leiicidius pectinijer and Siphatcles 
obesits. The example of 'ohesiis he figured shows 
about 22 rakers and presumably represents 
(1 ) hybrid origin; or (2) introgression from 'pectini- 
jer,' which occurred in the same lake (Winne- 
mucca ) ; or ( 3 ) an independent modification of 
G. b. obesa toward the lacustrine type. The two 
types both occur in some lakes with a complete 
range of intermediates, due apparently to hybrid- 
ization and intergradation. 

The marked differences in raker length, and 
correlated differences in their form, have been 
confirmed by casual examination of many more 
specimens than are entered in table 25. Such 
examination, for example, was accorded nearly 
all of the specimens, mostly of small size, that 
were utilized in enumerating the pharyngeal teeth 
(table 23). 

Although a positive correlation exists in gen- 
eral in Gila bicolor between the length and the 
number of gill-rakers — particularly exemplified 
in contrasting typical populations of G. b. obesa 
and G. b. pectinijer — no sharp or regular correla- 
tion exists among the stocks treated in this report. 
The adaptations of increased number and in- 
creased length of rakers have apparently been to 
a large degree independent. The rather complex 
relations between the mean length of longest 
raker ( table 25 ) and the mean numbers of rakers 
( table 22 ) become evident when these two param- 
eters are plotted together (fig. 29). The samples 
(1-4) from the Humboldt River system (fig. 
28A) have similar, median raker lengths, but de- 
crease regularly in mean number from down- 
stream to headwaters. In the two populations 
from Diamond Valley, regarded as variants of G. 
b. obesa, the rakers are alike in median number 



23 


~ 








q: 22 
g 21 


- 


•6 






_) 
_] 

5 20 










h 19 

z 
3 18 


- 


• 5 






° 17 

X 

i 16 

Ld 

_l 


• II 


• 3 

• 4 


•2 


• 1 


LJ 15 
> 

< 14 

UJ 

q: 

< 


- 


•10 
.7 






5 12 


" 


• e 






1 


1 


12 13 14 


15 16 


17 




MEAN NUMBER OF GILL 


RAKERS 





Figure 29. Correlation between mean number of 
gill-rakers (from table 22) and mean relative length of 
longest raker (from table 25), in various populations of 
Gila hkoltir in Nevada. The nimierals on the graph 
represent the Locations, Gl to Gil (listed on table 22). 

but are definitely longer than in typical 'obesa,' 
moderately so at Birch Ranch ( G5 ) and strikingly 
so ( fig. 28B ) at Sulphur Spring ( G6 ) . With some 
variation, both lengths and numbers are low in the 
subspecies G. b. lu'warlxcnsis (G7, 8; fig. 28C) and 
G. b. eucliila ( G 1 ) of the basin of Lake Newark. 
Median length and the lowest mean number char- 
acterize G. b. isolata (Gil) of Independence 
Valley. 

The form and texture of the rakers are also 
subject to marked differences, as is indicated es- 
pecially in the descriptions of G. b. riewarl<ensis 
(p. 168) and the Sulphur Spring race of G. b. 
obesa (p. 156). 

Pharyngeal teeth. In the vast area com- 
prising the many basins that were demonstrably or 
putativcly part of the watershed of Lake Lahontan 
at the height of the late pluvial period, or are 
thought somehow otherwise to have derived their 
Gila bicolor stock from the Lahontan drainage 
basin, the number of these teeth ( table 23 ) looms 
in importance. One reason is that the two main 
Lahontan forms, G. b. obesa and G. b. pectinijer. 



148 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 23. Niimhcr of pliaryni^cal Icclh in populations of Ciila hicolor //; vcitain basins in Nevada. 



Subspecies 

Pluvial lake system 
Locality 




No. 


on left side 


No. 


on ri 


ight side 




No. on either side 


4^ 


5 


6= 


No. 


Mean 


3^' 4 


5' 


No. 


Mean 


3 


4 


5 


6 


No. 


Mean 


Gila tricolor ohcsa 
































Lake Lahontan 
Carson River' 


-) 


35 


O 


39 


5.00 


— 25 


6'^ 


31 


4.19 


— 


27 


33 


-)^' 


62 


4.58 


Hiimhokit River 
Near Lovelock 
Near Carlin 

Bishop Creek 


3 
1 


40 
12 
14 


— 


42 
15 
15 


4.95 
4.80 
4.93 


— 38 

— 14 
1 14 


4 
1 


42 
15 
15 


4.10 
4.07 
3.93 


V 


40 

17 
15 


44 
13 
14 


— 


84 
30 
30 


4.52 
4.43 
4.43 


Lake Diamond 
Birch Ranch 
Sulphur Spring 


1 


14 
15 


— 


15 
15 


4.93 
5.00 


— 15 

— 15 


— 


15 
15 


4.00 
4.00 


— 


16 
15 


14 
15 


— 


30 
30 


4.47 
4.50 


Gila hicolor )icwaikcusis 
































Lake Newark 

Near Diamond Peak 
Moores Ranch 
Warm Springs 


3 


13 

12 

4 


— 


15 
15 

4 


4.87 
4.80 
5.00 


— 14 

— 13 

— 4 


1 
-> 


15 

15 

4 


4.07 
4.13 
4.00 


— 


16 

16 

4 


14 

14 

4 


— 


30 

30 

8 


4.47 
4.47 
4.50 


Gila hicolor euchila 
































Lake Newark 

Fish Creek Springs 




15 


_ 


15 


5.00 


— 15 


— 


15 


4.00 


— 


15 


15 


— 


30 


4.50 


Gila hicolor isohita 
































Lake Clover 

Independence Valley 


■> 


13 





1? 


4.87 


— 15 


— 


15 


4.00 


— 


17 


13 


— 


30 


4.43 


Totals 


16 


187 


T 


205 


4.93 


1 182 


14 


197 


4.07 


1 


198 


193 


2 


394 


4.50 



differ ill tlii.s respect (as well as in the numbers of 
gill-rakers and pelvic rays, and in various features 
of form ) . 

Gila hicolor ohesii (pp. 153. 156). the more 
widespread type, typically inhabiting streams and 
springs, normally has 5 — 4 teeth ( that is. 5 on 
the left arch and 4 on the right ) . and this is true 
also of the modified derivatives of G. h. ohcsa 
in the enclosed basins just south of the Humboldt 
River system (table 1). Rarely the numbers are 
reversed. 4 — 5. or the same number occurs on 
both sides. When symmetrical in number, the 
teeth are more often 4 — 4 than 5 — 5, except 
where, as in the samples from Carson River near 
Fallon and from Humboldt River near Love- 
lock, introgression from 'pcclinifcr' has ap- 
parently increased the proportion with 5 — 5 teeth. 
In the sample from Bishop Creek we have found 
a variant with 4 — 3 teeth. Reduction to 3 teeth, 
in this case on the left arch (formula 3 — 4) was 
found (along with one of 4 — 4) among 10 counts 
for the introduced stock of Gila hicolor ohcsa in 
Little Soda Lake. Nevada. 

In contrast, the teeth normally number 5 — 5 in 



the lacustrine form, which Snyder (1917, pp. 
64-67, figs. 5, 6) regarded as generically as well 
as specifically distinct, under the name Lcucidius 
pcctinifcr. but which, by reason of evidence of 
extensive introgression and intergradation (pp. 
143-149). vve separate only subspecifically. as 
Gila hicolor pcctinifcr. Snyder stated that the 
teeth in this form are invariably 5 — 5. but we find 
some with 5 — 1. 

Precautions in the counting of the pharyngeal 
teeth when some have been lost are mentioned on 
page 93). 

Subspecies status of isolated forms. We 
combine within one species, Gila hicolor. not only 
the ecologically segregated forms discussed above 
as G. h. ohcsa and G. h. pcctinifcr. but also all 
of the hitherto described, isolated local forms, 
including those of the Death Valley system ( Mil- 
ler, 1973). that have been referred to Siphatcles 
( and may be retained in a subgenus of that name). 
We have proposed three additional subspecies. 

All of these forms, plus a considerable number 
that remain to be named, have many features in 
common, though they differ greatly in the number 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



149 



Table 23. Continued. 



Subspecies 
























Asymmetry' 




Total number of teeth 




Tooth formulas" 




100(L+R) 




Pluvial lake system 
Locality 
























IGOR 


T 


8 


9 


10 


No. 


Mean 


4-3' 4-4 


4>_5' 


5-4 


5-5* 


6=-4 


No. 


L+R 


Gila bicolor obesa 




























Lake Lahontan 




























Carson River^ 


— 


— 


25 


e 


31 


9.19 


— — 


-) 


23 


4^' 





87 


7 


Humboldt River 




























Near Lovelock 


— 


T 


36 


4 


42 


9.05 


"1 


— 


36 


4 


— 


90 





Near Carlin 


— 


3 


11 


1 


15 


8.87 


— 3 


— 


11 


1 


— . 


73 





Bishop Creek 


1 


— 


14 


— 


15 


8.87 


1' — 


— 


14 


— 


— 


100 





Lake Diamond 




























Birch Ranch 


— 


1 


14 


— 


15 


8.93 


— 1 


— 


14 


— 


— 


93 





Sulphur Spring 


— 


— 


15 


— 


15 


9.00 


— — 


— 


15 


— 


— 


100 





Gila bicolor ucwarkcnsis 




























Lake Newark 




























Near Diamond Peak 


— 


T 


12 


1 


15 


8.93 





— 


12 


1 


— 


80 





Moores Ranch 


— 


-> 


12 


1 


15 


8.93 


-> 


1 


11 


1 


— 


80 





Warm Springs 


— 





4 


— 


4 


9.00 


— — 


— 


4 


— 


— 


100 





Gi}a bicolor ciichila 




























Lake Newark 




























Fish Creek Springs 


— 





15 


— 


15 


9.00 


— — 


— 


15 


— 


— 


100 





Gila bicolor isolata 




























Lake Clover 




























Independence Valley 


— 


-» 


13 


— 


15 


8.87 


— 2 


— 


13 


— 


— 


80 





Totals 


1 


12 


171 


13 


197 


8.99 


1 12 


3 


168 


11 


2 


88 


T 



1 When the number of teeth is reduced to 4 on the left side, tooth no. 1 (uppermost) is usually more or less disproportionately massive. 

- In the two specimens with 6 teeth in the major row on the left arch the seriation of the teeth is slightly irregular, as is indicated in the text. 

■'In the specimen with 4 — 3 teeth there is evidence (see text) that there was not quite enough room for the uppermost tooth to fit. 

' When 5 instead of 4 teeth develop on the right side, and soiTietimes when 5 form on the left arch, the uppermost tooth is more or less reduced 
and displaced outward, more or less against the side of the second tooth. 

^ The population from the Carson River near Fallon represents G. b. obesa modified by introgression from G. b. pectinifer. This is reflected by 
the increased number of fish with 5 teeth on each side (a characteristic of G. b. peclinifer, but otherwise essentially like the other specimens in 
the scries. 

"In the tooth formula the left side is represented first. 

'The indices, as proposed by Hubbs and Hubbs (1945, pp. 2.11-233), specify (1) the degree of .asymmetry in the count and (2) among asymmetri- 
cal counts, the percentage of extrality (normally the count on the lett side is c>ne higher than on the right side). 



of gill-rakers. No specimen in the entire comple.x 
has ever been found to have teeth in other than 
the main row. In all, the better developed scales 
have a distinctive form: shield-shaped, with 
focus far basad, typically with apical radii only, 
circuli well spaced in the apical field, moderately 
crowded laterally, and densely crowded basally; 
with strong anterolateral angles, and with the an- 
terior margin and anterior circuli arched inward 
toward each anterolateral angle ( as illustrated on 
fig. 46A and by Kimsey. 1954. fig. 4); some 
scales, however, are more nearly vertically oval, 
without sharp angulation. As a rule, the various 
local forms are moderately large and have rela- 
tively uniform contours, proportions, and colors 
(figs. 30, 31, 35A,B). The range of the species. 



though in arid areas rather scattered, is relatively 
compact. 

When considered alone, some of the localized 
forms seem sufficiently distinctive to lead one to 
question whether full specific separation would 
not be proper, but when the whole array of forms 
is kept in mind, subspecies status seems preferable. 
No two forms of the complex, other than 'obcsa 
and 'pectinifer,' are anywhere sympatric. 

Local Populations Of Gila bicolor 
Examined 

The populations of Gila bicolor treated in de- 
tail in the present study, all in Nevada, are re- 
ferred to four subspecies ( listed also, briefly, in 
table 21): 



150 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 24. Conclalinn of i;ill-nikcr nunihcr with lite tonlli ftutlcin.s (harmlerisric of Gila bicolor obesa (5-4) and 
of G. b. pectinifer (5-5) for mo Locations in the Lahontan system. 



Location Gl. Carson River near Fallon (UMMZ 124859) 



Tooth 
formula 



Gill-raker number (both sides counted) 



12 



13 



14 



15 



16 



17 



n 



19 



20 



21 



23 No. 



(rarely 4 — 5) 



5—5 



10 10 
— 1 



50 



Ave. 



16.72 

17.75 



Location G2. Humboldt River near Lovelock (UMMZ 124873, 124874) 

Gill-raker number (both sides counted) 

Tooth 

formula 12-16 17 18 19 20 21 22 No. 



Ave.' 



5—5 



64 



— 1 

4 — 



2 1 

— 1 



1 



72 
8 



15.46 
18.13 



'Average computed on assumption that the raker number tor the entire 12-16 mm. grouping was the modal number, 15 (table 22). 



(A) G. h. ohesa (Girard), represented by samples from 

the following Locations: 
Gl: Carson River near Fallon, near central part 

of Lake Lahontan drainage basin. 
G2: Humboldt River near Lovelock, on the bed of 

I akc Lahontan. 
G3: Humboldt River near Carlin, above the bed 

of Lake Lahontan. 
G4: Bishop Creek, a headwater of Humboldt River. 

near Wells. 

(A') G. h. ohcsa. aberrant races: 

Cj5: Birch Ranch, on east side of Diamond Valley. 
G6: Sulphur Spring, on west side of Diamond 
Valley. 

(B) G. h. tu-warkensis Hubbs and Miller, represented by 

samples from three Locations on the bed of 
pluvial Lake Newark: 
G7: Spring near Diamond Peak, on west side of 

Newark Valley. 
GS: Spring-led pool at Moores Ranch, on west 

side of Newark Valley. 
G9: Warm Springs, near north end of Newark 
Valley. 

(C) G. />. ciultila Hiibhs and Miller, from a single Loca- 

tion, also in the basin of Lake Newark: 
GU) and lOA: Fish Creek .Springs, in L'lsh Creek 
Valley. 

(D) G. /'. isoliiia Hubbs and Miller, represented from a 

single Location in the basin of pluvial Lake 
Clover: 
Gil: Warm Springs, Independence Valley. 

On the several maps (figs. 3, H. 12) involved, 
the Locations (the extraiiniital Gl and Cj2 ex- 



cepted) are marked, in the sequence of the pre- 
ceding enumeration, by the designations G3 to 
GIL^ 

The seemingly remote possibility that some form 
of Gila bicolor ( it would most plausibly have 
been G. h. obesa) occurred until recently in 
Ruby Lake of the Lake Franklin basin is suggested 
by local testimony (p. 209). 

Lahontan Creek Tui Chub 

Gila bicolor obesa ( Girard ) . 
(Figure 30.) 

As mentioned in the preceding account of the 
species, we treat the trophically differentiated 
stream and lake forms of the subgenus Sipluiteles 
in the Lahontan .sy.stem and related drainage 
basins, namely 'obesa and 'pccliiiifer,' as sub- 
specifically rather than as specifically distinct (or 
as taxonomicaily inseparable). Our study of the 
entire complex has further suggested that the 
stream form of the Lahontan system is subspecifi- 
cally separable from Gila hicclcr bicolor of 
Klamath Lake, Oregon. 

Because the populations of Gila bicolor that 
occur in isolated waters within the area of Nevada 
included in this report all give evidence of having 
been derived from stocks of G. b. obesa (or an 
imntediate ancestor) of the Humboldt River sys- 
tem, we iiave included in our analysis, as we did 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELIC T FISHES 



151 



for Rhinichthys osciiliis, samples from that system 
taken to represent the form ancestral to the now 
isolated stocks. Location G2 is from near Love- 
lock, close to the lower end of the river, on the 
bed of Lake Lahontan, probably from near where 
the types were collected. Location G3 is from 
midcoiirse of the Humboldt River, near Carlin. 
Location G4 is from a headwater tributary. Bishop 
Creek. We have incorporated also data for a 
collection (Gl) from the Carson River near 
Fallon, also on the bed of Lake Lahontan, to 
represent a stock of G. h. ohcsa that appears to 
have been modified by introgression from the 
lacustrine form, G. h. pcctinifi'r. 

Populations of Carson River and Humboldt 
River Systems. 

Location G I .— Clarsoii River, in T. 19 N., R. 28 E.: 
Churchill County. 2 miles west of Fallon, Nevada, on 
the bed of Lake Lahontan (west of the limits of fig. 1 ). 
Rather muddy water (bottom visibility about 0.3 m.); 
mud and gravel bottom; pools and riffles; scanty growth 
of MyriophyUum and Potainogeton; 20" C. (air .31°). 
Stream here 7-20 m. wide and to 0.6 m. deep. Hubbs 
family and Miller, July 30, 193,S (M3S-77); UMMZ 
1248.*59 and 133244 (35, 28-136 mm.); 2.5-1001 seine 
with '4 -inch square mesh in bag. Associated species 
were four other natives (Catostonuis tahoensis. C. platy- 
rliyuchus. Rhinichthys osciilii.s rol'usius. Richanlsoiiiiis 
egrcgiiis) and four exotics {htaliiiiis in. nicliis. Pen a 
flavescens. Micropterns snlnmidcs. and Anhoplites in- 
terriiptns ) . 

Location G2. — Hiinihoklt River near Lovclotk, at 
Irish American Dam, in T. 27 N., near boundary of 
R. 31-32 E.; Pershing County, Nevada, about 3 miles 
northeast of Lovelock, on the bed of Lake Lahontan 
(west of fig. I ). Water olive-yellow and muddy (bottom 
visibility ca. 15 cm.); mostly firm to soft clay, with fine 
gravel in riffles; pool-and-riffle; scanty growth of Pota- 
niogeton, cf. P. pectinatus; 23° C. (air 32°). Stream 
here 5-20 m. wide and to ca. 1 m. deep. Hubbs family 
and Miller, July 31. 1938 (M38-79); UMMZ 124873-74 
(111, 20-115 mm.); 25-foot seine with '4 -inch square 
mesh in bag. Some of the young and half-grown from 
the backwater had mud on gills. Four other native 
fishes, Caloslonuis tahoensis. C. plutyrhynchus (as well 
as hybrids between the two suckers), Rhinichthys uscnius 
robustus. and Richardsonius egregius. and three exotic 
species (Cyprinns carpio. hialuiiis n. nelniloMis. and /. 
nt. nu'lds) were collected. 



Location G3. — Humboldt River near Carlin, just 
above hot springs along and in river bed, in T. 33 N., 
R. 52 E.; Elko County. Nevada, ca. 1.0 mile southwest 
of Carlin (figs. I. 8). Rather murky (clay limiting bot- 
tom underwater visibility to ca. 0.5 m.); clay, sand, and 
mud. with gravel on riffle; pool, riffle, and backwater 
slough; no vegetation; 23° C. (air 33°). Hubbs family 
and Miller, August 11, 1938 (M38-119); UMMZ 124915 
(40. 27-107 mm.); 25-foot seine with '4 -inch square 
mesh in bag. Associated species were four other natives 
(Catostonuis tahoensis. C. plutyrhynchns. Rhinichthys 
oscidns robnstiis, Richardsonius egregius ) and three 
exotics (Cyprinns carpio, Ictalnrus n. nehidosus, I. in. 
inelas ) . 

Location G4. — Bishop Creek: described (p. 107) 
under Location R2 (fig. 12); UMMZ 141523 (254. 17- 
137 mm.). Young to adult abounded. The young 
swarmed along margins of pond. Some of the females, 
taken off a large gravel riffle, were running-ripe. Three 
other native fishes were obtained (Catostonuis tahoen- 
sis. Rhinichthys oscidiis robustus. Richardsonius egre- 
gius ) . 

The Lahontan populations sampled, one from 
Carson River and three from the Humboldt River 
system, are by no means uniform in systematic 
characters. In most respects the characters form 
a downstream-upstream dine. In some respects, 
notably gill-raker number, the ciine in the down- 
stream direction verges toward Gila bicolor pccti- 
nifcr. The ciine in the upstream direction ap- 
proaches the forms that inhabit the endorheic 
basins just south of the Humboldt River. 

In the following analyses the variation, 
whether or not clinal, is treated in the upstream 
direction, taking the Carson River sample first 
(Gl ^G2-G3-G4). 

Size. The maximum standard lengths in this 
.series are 1.36 ^ I 15 — ' 107 ^ 137 mm. Rather 
contrary to expectation, no significant trend is 
indicated. 

Coloration. The coloration is essentially 
plain and, as the species name implies, bicolored: 
pigmented on the back and midsides and silvery 
below (figs. 30A-C). The bicolored condition 
and the general light tone is more conspicuous in 
the downstream Locations (Gl and G2); in the 
two upstream Locations ( G3 and G4 ) the fish are 
duskier and the puncticulate area extends farther 



152 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



down, leaving little but the transverse ventral 
region silvery. In the series as outlined, there is 
a gradation in the pigmentation along the middle 
of the trunk, from a pattern essentially margining 
the silvery centered, vertically diamond-shaped 
scale pockets to one more evenly darkened. There 
in a striking upstream gradient in the shape and 
intensity of the basicaudal spot, which is higher 
than long, pointed forward, and truncate behind: 
it is strong in the Carson River sample, somewhat 
weaker and more diffuse in the collection from 
near Lovelock, weak to diffuse or small in the 
Carlin lot. and barely evident in the Bishop Creek 
collection. 

In all series, the head is dark on top, on the 
snout, around the lower border of the eye, and 
on the upper part of the opercular region; large 
melanophores are conspicuous on the cheek, the 
opercular region, and the lower sides. In all 
series, the dorsal and caudal fins are dusky and in 
nearly all specimens there is some puncticulation 
on all lower fins. 

As in all forms of Gila hicolor here treated, a 
rather diffuse underlying midlateral dark streak 
is developed in young fish, but tends to become 
obscure or obsolete in adults; the young also show 
scattered large melanophores on the lower sides — 
a distinctive feature of the whole species. 

Life colors. The field notes indicate that at 
least one of the specimens taken near Carlin had 
the center of the lower fins rusty orange, and 
that the males collected at Bishop had some gilt 
on the sides and the lower fins brownish tan inside 
light margins. 

Form. The general form ( figs. 30A-C ) grades 
in the stated series from one that downstream is 
rather strongly compressed and sharp-snouted, to 
one upstream that is more chubby and turgid, 
with a somewhat rounder snout in both side and 
top view. The fins grade, in the same direction, 
from slightly falcate to moderately rounded. 

Lateral-line system. Ordinarily, the lateral 
line on the body is complete in G. b. obesti. 

The supratemporal canal (table 26) is more 
often incomplete than complete in these four col- 



lections, but only slightly so at Location G2 near 
Lovelock. 

Morphometry. For all three Locations in the 
Humboldt River .system, the morphometric data 
( table 27 ) were compared between smaller and 
larger specimens of each sex, within standard- 
length groupings, of 45-98 and 100-137 mm. 
for females and of 48-73 and 74-107 mm. for 
males (one 76-mm. male from Bishop Creek was 
included in the smaller category, because no 
others in the large category are available). Within 
the downstream-to-upstream series the propor- 
tional measurements show for each sex an in- 
crease in the predorsal length, in line with a trend 
toward a posterior position of the dorsal fin in 
isolated waters, a slight increase in head size, and, 
in line with expectation, a definite decrease in 
average size of fins. 

The mean values for proportional predorsal 
length (fig. 36) for the Humboldt River samples 
are relatively lower than in the isolated subspecies, 
and the mandible (figs. 34, 37) averages shorter 
than in G. b. ciichihi and G. b. isolata. 

Sexual dimorphism and nuptial charac- 
ti:rs. In the three series from the Humboldt 
River system the sexual dimorphism is rather 
limited, though in the usual direction (table 28). 
The dorsal fin usually averages less pcisterior in 
males, but the difference is small except for the 
larger fish from near Lovelock. The enlargement 
of the paired fins in the males is about average: 
the excess of the means in the males, in terms of 
percentage of the standard length, is 0.1-1.6 for 
the dorsal, 0.4-0.8 with one obviously unrepre- 
sentative figure of 4.4 for the caudal. 2.2-3.4 for 
the pectoral, and 0.8-1.6 for the pelvic. The 
moderate size of the pectoral fin in the males of 
G. b. obesa is emphasized by comparing the values 
for this subspecies with those of G. b. newaikensis 
(fig. 32). 

A 76-mm. nuptial male from Bishop Creek dis- 
plays nuptial tubercles similar to those described 
forC. b. isolata (p. 179). 

Fin rays. Rays were counted (tables 29, 30) 
on all four series from the present Lahontan drain- 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



153 



age system. In the entire lot, the dorsal-ray counts 
seldom vary from 8. The number of anal rays 
decreases slightly on the average upstream to Car- 
lin. in sHght approach toward the number, 
modally 7, in the subspecies confined to isolated 
springs, but seems to revert to a slightly higher 
average in Bishop Creek. The caudal rays vary 
from 17 to 20, with few variants from 19. The 
pectoral-ray mean decreases slightly in the up- 
stream gradient within the Humboldt River sys- 
tem, but is about average in Carson River. The 
pelvic-ray mean fluctuates sHghtly: 9.49 near 
Lovelock, 9.29 near Carlin and in Bishop Creek, 
and only 9.21 in Carson River (which is sur- 
prising, because this series showed signs of intro- 
gression from G. b. pectinifcr. which usually has 
10 pelvic rays). 

Vertebrae. The vertebral counts ( table 3 1 ) 
fluctuate shghtly and irregularly. 

Scale rows. The average for the 1 1 scale 
counts (table 32) is uniformly highest in Carson 
River, which may reflect the higher number in 
G. b. pectinifer. The sequence in the downstream- 
upstream series (Gl to G4) was either 2-^4—' 
3-1 or 4^2—3-^1. 

Gill-rakers. The raker counts (table 22) 
exhibit a very sharp upstream decrease, in a 
regular cline, with means of 16.43 > 14.90 > 
13.62 > 12.61. As noted above, we attribute the 
higher number downstream to past hybridization 
with G. b. pectinifer. which has 29 to 40 rakers, 
and subsequent backcrossing with G. b. obesa. 
The strong positive skewness of the counts for 
Carson River and for Humboldt River at Love- 
lock, with a long gradual tail in the distribution, 
is a major consideration in our interpretation. 
Especially notable is the lack of positive skewness 
for the two upstream collections, far removed 
from contamination with G. b. pectinifer. The 
finding of intergrades between G. b. obesa and 
G. b. pectinifer in ancient fish caches of Indians 
in the Lovelock region, as we have pointed out 
(Hubbs and Miller, 1948b, p. 41 ), demonstrates 
past intermixing near the lower end of Humboldt 
River. The collections from Carson River near 



Fallon and from Humboldt River near Lovelock 
appear to represent largely reintegrated popula- 
tions, rather than a mixture of G. b. obesa with Fi 
hybrids between G. b. obesa and G. b. pectinifer. 
The extreme upper end of the counts in the two 
series is about midway between the mean for G. b. 
obesa and the midpoint between the means for the 
two forms, and the close similarity to G. b. obesa 
of the individuals with high raker counts suggests 
to us past backcrossing with G. b. obesa. the 
locally dominant form. 

The gill-rakers in the Humboldt River system 
are of moderate length, shorter than in the Dia- 
mond Valley populations of G. b. obesa and 
longer than in G. b. newarkensis and G. b. eiicliila 
(table 25; fig. 28). 

Pharyngeal teeth. Influence of G. b. 
pectinifer in the collections from Carson River 
near Fallon and from Humboldt River near Love- 
lock seems to be indicated also by the somewhat 
higher than usual proportion of pharyngeal 
arches with the 5 — 5 tooth formula of the lake 
form — more frequently 5 — 5 in these collections 
than in any other populations here treated ( table 
23). That the specimens with 5 — 5 teeth have 
almost the same raker number as those with 5 — 4 
teeth in the Carson River sample and a not much 
higher raker number in the sample from near 
Lovelock, however, is shown by correlating the 
tooth and raker counts ( table 24 ) . The dental 
formula in these samples, especially the one from 
Carson River, is also more variable than in other 
samples, which in this regard are essentially uni- 
form. 

Aberrant Populations, of the Pluvial Lake 
Diamond Basin, Referred to Gila bicolor 
obesa. 

The two populations in this category occur 
within the sump of the extensive drainage basin of 
pluvial Lake Diamond, close to the margin on 
the ancient lake, one on the west side and one on 
the east side. Field reconnaissance, with frequent 
interviews, yielded no seemingly reliable indica- 
tion that tui chubs (Gila bicolor) occur in the 



154 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 25. Li'n,i;lli of longest gill-nikcr in populations of Gila bicolor in certain basins in Nevada. 



Subspecies 

Pluvial lake system 
Locality' 



Gitu bicolor ohcsa 
Lake Lahontan 
Carson River ( I ) 
HunihokIt River 
Near Lovelock ( 2 ) 
Near Carlin (3) 
Bishop Creek (4) 
Lake Diamond 
Birch Ranch (5) 
Sulphur Spring (6) 
(lila bicolor iic\rarkcnsis 
Lake Newark 

Near Diamond Peak ( 7 I 
Moores Ranch (8) 
Gilii bicolor ciichihi 
Lake Newark 

Fish Creek Springs (10) 

Gila hicolor isohaa 
Lake Clover 

Independence Valley (11) 



Length of rakers' in permillage of standard length' 



7.5-11.5 11.5-15.5 15.5-19.5 19.5-23.5 23.5-27.5 27.5-31.5 



I 


14 


16 


3 


— 




10 


41 


3 


— 


— 


8 


35 


3 


— 


— 


11 


24 


3 


— 




4 


11 


10 


1 


— 


2 


9 


25 


IS 


y 


37 


■6 






16 


23 


— 


— 


— 



1 



30 



10 



2<i 



40 



' Measured on one side from liard base to tip. 

'- Expressed as numbered Locations in tlie G series. 

■Values grouped in classes of four units; mean values of units are lislcU. 

' Means delerniincd from ungrouped data. 



N 



34 



58 



56 



M* 



15.8 



54 


16.7 


46 


17.0 


38 


16.6 


26 


19.0 


57 


22.1 


54 


13.4 


39 


12.1 



14.9 



16.9 



tributary basins to the west and southwest, where 
two populations ot Rhiniclithys oscuhis were dis- 
covered. Our field woriv convinced us that on the 
east side of Diannmd Valley chubs occurred tmly 
at Birch Ranch (Location G5). along with 
Rhinicluliys osciiliis (p. 115). On the west side 
we found them only in Sulphur Spring (G6, where 
it alone occurred ). and deteniiincd it to be lacking 
in Big Shipley Spring, which harbored Rltiitichlhys 
(Location R(S). No definite indication was ob- 
tained, by local inquiry or examination, of the 
occurrence of Gila hicolor elsewhere on the west 
side of Diamond Valley, but the possibility re- 
mains that some may have held out in some of 
the springs we did not examine (pp. IS-19). 
We did hear one vague report of fish called 
"chubs' in a spring hole in the yard of Bailey 
Ranch .^v2 km. south of Big Shipley Spring. We 
think it probable that only the two poptilations, 
or at least types, of Gila bicolor persist (or oc- 
curred in I'^J.iS) anywhere in the wide arid ex- 
panse of the ancient Lake Diamond drainage 
basin of Nevada. 



Locatioti G5. — Springs on Birth Rantli: described 
(p. 115) under Location R7 (figs. .3. S). UMMZ I 24'^)34 
(113, 15-Nl nun.l. The prestiniabK indigenous occur- 
rence ol this chtib here, and its close association with 
Rhini( htlivs oscnliis. are discussed on the same page. 

Location Cid. — .Sulphur Spriiiu;, on the west side of 
the extensive Hat of Diamond Vallcs. just below the 
loot of the Sulphur Spring Range, shown on the Garden 
Valley 15-nunule Quadrangle as near the western edge, 
submcdially, ol ,Sec. 36, F. 23 N.. R. 52 l-.; eastern 
Eureka County. Nevada, .S.7 nnlcs by road south of 
Sadler Ranch (tigs. 3, S). Somewhat stilphurous, and 
clear btit casih' roiled; very soft pulpy peat; no current; 
much algae, rushes, etc.; 22 C. (air 2S' ). Htibhs family 
and Miller, August 12, 1038 (M3S-I22): UMMZ 
124'-»27-2S (71. 2I-S(i mm.); 0-foot woven-mesh seine 
•ind 15-foot seine with 'j-inch square mesh. 

The Sulphur Spring population .seems to be 
rigidly confined to the somewhat dammed spring 
pool which was about 10 m. in diameter and 
about 0.3 111. deep to an ill-defined bottom. No 
fish were in the irrigation ditch leading out from 
the pool. Algae were observed in the feces of the 
fish. 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



155 



In consonance with the evidence of a rather 
late-pluvial outlet of Lake Diamond ( pp. 1 6- 
17). the two populations of Diamond Valley 
are referred to the Lahontan creek tui chub, Gila 
bicolor obesa. However, the Sulphur Spring pop- 
ulation (G6) would have been separated sub- 
specifically, chiefly on the basis of its unusually 
long gill-rakers (table 25; fig. 28B), had we not 
found that the Birch Ranch series (G5) bridges 
the gap. Differences between the two isolated 
spring populations of Diamond Valley, as speci- 
fied below, indicate considerable modification of 
the Diamond Valley stock since Lake Diamond 
ceased discharging in presumably rather late 
pluvial time, and also suggest some differentiation 
since Lake Diamond desiccated beyond conditions 
viable for fish, probably only a few thousand 
years ago. 

Size. Consistent with the highly restricted 
habitat, the two populations are rather dwarfed. 
The largest specimens, as now preserved, are 81 
and 86 mm. in standard length, respectively, as 
compared with 107-137 mm. for the samples 
from Carson and Humboldt rivers. 

Coloration. The general coloration (fig. 
30D) in both lots is essentially the same as in the 
upstream samples of the Humboldt River system. 
The basicaudal spot varies from moderately strong 
to minute and/or diffuse, or even lacking. It is 
apparently never vertically elongate, with a 
definite anterior angle and a sharply truncate pos- 
terior edge, as it is in the two downstream popula- 
tion discussed above. 

Life colors. The colors of the Sulphur Spring 
fish were described as follows in the field, and 
it was noted that those from Birch Ranch were 
similar: rather bright olive on back, passing 
through more or less definite gilt reflections on 
midsides to bluish white on lower parts. Dorsal 
and caudal fins olive. Lower fins olive, grading 
toward yellowish, with only a trace of a bluish 
border on pelvic and anal. Axil of pectoral fin 
and exposed part of shoulder girdle more or less 
golden, with a pinkish tinge in some. Iris bright 
gold around pupil. These colors seem to have 



been somewhat modified from those displayed by 
the tui chubs in the Humboldt River and its tribu- 
taries. 

Form. The Birch Ranch specimens are shaped 
much like those from Bishop Creek, with moder- 
ately turgid body contours. The Sulphur Spring 
fish (fig. 3()D) tend to be rather more compressed, 
to have the dorsal contour more elevated, the 
muzzle more pointed, and the mouth more oblique 
and straighter ( which may be correlated with the 
extreme development of the gill-rakers). The fins 
are about as rounded as in the upstream end of 
the cline in the Humboldt River system. 

Lateral-line system. Ordinarily, the lateral 
line on the body is complete in each population in 
Diamond Valley. 

The supratemporal canal (table 26) seems to 
be uniformly incomplete in both collections from 
Diamond Valley, as in no other area treated in 
this study. This circumstance lends support to the 
view that the two populations have had a common 
origin. In many of the specimens, the canal has 

Table 26. Condition of supratemporal canal Iconi- 
pletc or incomplete) in populations of Gila bicolor in 
certain hasins in Nevada. 



Subspecies 


Supratemporal canal 


Pluvial lake system 
Locality 






Complete 


Incomplete 


Gila hiicolor ohcsn 






Lake Lahonlan 






Carson River 


6 


24' 


Humholdl River 






Near Lovelock 


20 


28 


Near Carlin 


6 


24= 


Bishop Creek 


12 


26 


Lake Diamond 






Birch Ranch 


— 


29= 


Sulphur Spring 


— 


35= 


Gila bicolor ncwarkciisis 






Lake Newark 






Near Diamond Peak 


24 


6 


Moores Ranch 


21 


9 


Gila bicolor euchila 






Lake Newark 






Fish Creek Springs 


17 


6 


Gila bicolor isolata 






Lake Clover 






Independence Valley 


27 


— 



' One break in 14. 2 breaks in 6, and 3 breaks in 4. 
-'One break in 12. 2 breaks in 5. and 3 breaks in 7. 
^ In many specimens the canal has 3 breaks, one at midline and 
one on each side. 



156 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



llirce breaks, one at the midline and one on eaeh 
side. 

Morphometry. Distinctive features in body, 
head, or fin proportions ( table 27 ) are few. The 
body seems to average deeper in the Diamond 
Valley series than in those from Humboldt River. 
The head in the Sulphur Spring specimens, but 
not in those from Birch Ranch, averages longer 
than in the Carson-Humboldt samples. The fins 
in the Sulphur Spring fish average somewhat 
larger than in those from Birch Ranch and defi- 
nitely larger than in the more upstream Carson- 
Humboldt samples. 

SixuAL DIMORPHISM. Aliiiost uo .sexual di- 
morphism is indicated for the body, head, and 
vertical-fin measurements (table 28). For pec- 
toral-fin length the excess values for the males, 
expressed as percentage of the standard length, 
are 2.6 and I.S for the pectoral fin for the Birch 
Ranch and Sulphur Spring collections, respec- 
tively. Corresponding values for the pelvics are 
1.4 and 0.9. These values compare closely with 
those for the present LahiMitan drainage. The 
enlargement of the fins in the male in these series 
is indicated by the measurements to be small 
for the dorsal fin, dubious for the caudal, and 
about average for the paired fins. 

Fin rans. The ray counts for the vertical fins 
(table 29) are not distinctive. The counts of 
pectoral rays (table 30) average very slightly 
lower in the Sulphur Spring than in the Birch 
Ranch sample, but fit within the variation for the 
Humboldt samples. The average for the pelvic- 
ray counts is very slightly reduced in the Sulphur 
Spring fish, as contrasted with the other samples 
referred to G. h. ohesa. 

VHRTiiBRAi:. Not distinctive (table 31 ). 

Scale rows. The scale-count averages ( table 
32) for all I I sets are virtually identical for the 
two Diamond Valley samples, and generally fit 
within the variation for the samples more typical 
of G. /'. ohcMi. The counts of predorsal scales 
and of scales around body average slightly lower 
than for the Humboldt River Locations. 

GiLL-RAKiZRS. The raker counts (table 22) 
are very slightly higher on the average in the 



Sulphur Spring than in the Birch Ranch sample, 
and correspond closely with the mean for the 
Carlin sample, in the .sharp dine of the Carson- 
Humboldt series. The averages are definitely 
lower than in the Lovelock downstream sample 
and are distinctly higher than in the Bishop Creek 
headwater sample. 

As already mentioned, the rakers in the Sulphur 
Spring sample are so outstandingly long (table 25; 
fig. 2KB) that a subspecies would have been 
erected for it, had not the Birch Ranch fish been 
so definitely intermediate in this respect between 
the Sulphur Spring sample and all others referred 
to Gila hicolor ohcsci: 81 percent of the raker 
measurements for the Sulphur Spring sample, ex- 
pressed as permillage of the standard length of the 
fish, are barely overlapped by less than 10 per- 
cent of the values for any of the four samples 
from the Carson and Humboldt localities, but in 
this respect the Birch Ranch fish are just inter- 
mediate. 

The gill-rakers of the Sulphur Spring tui chubs 
also differ in form and structure (fig. 28B) from 
those of approximately topotypic G. b. obcsa, as 
represented by the Lovelock series. They remain 
bony and slender to their well separated bases, 
and there are usually some crenulations along the 
inner edge. In typical G. /'. ohcsd the rakers 
(fig. 28A), especially in small specimens, grade 
in appearance from those of G. h. iicwurkcnsis 
( p. I 68; fig. 28C ) to those of the Sulphur Spring 
sample (fig. 28B). 

Pharyngeal teeth. The dentition of the Dia- 
mond Valley populations is essentially like that 
of G. h. ohc'sci. Among 15 specimens examined 
for each population the only variant from 5 — 1 
is one from Birch Ranch with 4 1. 

Newark Valley Tui Chub 

Gila bicolor nevvarkensis Hubbs and Miller. 
(Figure -^lA.B.) 

Clild hicolor ncwtirkcnsis HltHliS and Mu LER, l'J72, p. 
102 (diagnosis). 

All indications point to the conclusion that 
this subspecies is confined to the main depression. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



157 



Newark Valley proper, in the drainage basin of 
pluvial Lake Newark (pp. 22-26; figs. 3, 8), in 
the western part of White Pine County, Nevada, 
and that it is the only native fish in the main 
depression. The form occurring in Fish Creek 
Springs, in Fish Creek ( Little Smoky ) Valley, 
the southwestern arm of the same drainage basin, 
is regarded as constituting a differentiated sub- 
species, Gila bicolor cuchila (pp. 168-174). 
Populations of G. b. newarkcnsis have been taken 
in only three springs, with data as follows: 

Location G7. — Spring in Newark Valley on west side, 
near Diamond Peak (called "South Peak" locally in 
1934), on alluvial slope about opposite south end of 
Newark Dry Lake, now thought to he at or near Circle 
Ranch as shown on the Eureka 15-minute Quadrangle 
and as Labarry Ranch on a county map of 1959. or at 
one of the springs mapped for about 2 km. south and 
southwest of that ranch, near middle of T. 20 N.. R. 
55 E.: in northwestern White Pine County, Nevada 
(figs. 3, 8). Clear water: soft bottom (person sank 
about 15 cm.); seepage overflow; Cliani forming a thick 
mat on bottom, algae and some NciMiirfiiini blanketing 
surface; water cool; pool ca. 7x12 m. and ca. (1.6 m. 
deep. Hubbs family, September 11, 1934 (M34-206); 
UMMZ 132185 and 188893 (227, 25-97 mm.); 15-foot 
seine with 'a -inch square mesh. Some extremely young 
fish seen here (and at GS) indicated late spawning in 
the cold water. 

Location G8. — Spring-fed pool on west side of New- 
ark Valley, on Moorcs Ranch, apparently the one 
named Goecoechia Ranch on the Diamond Springs 15- 
minute Quadrangle, and probably one of the pools 
shown between the 5,860-foot and 5,880-foot contours 
in SW I4 Sec. 11 and NW '4 Sec. 14, T. 22 N., R. 55 
E., in northwestern White Pine County. Nevada, by road 
6 miles north of Strawberry (Strawberry Ranch) and 5 
miles south of Simonsen, also known as Simonsen Ranch 
or Cold Creek Ranch (figs. 3, 8). Clear water; soft 
mud; virtually no current; only a few open spots in 
dense Nastiirtiuin, with Cliara thick below; water cold. 
Hubbs family, September 11, 1934 ( M 34-207 ) ; UMMZ 
132186 (113, 20-67 mm.); 15-foot seine with 14-inch 
square mesh. 

Location G9. — Ditch fed by Warm Springs, on what 
was known as "Billy Moore's Ranch," on the very gentle 
alluvial slope in the eastern lobe at the north end of 
Newark Valley, apparently between the 5,880-foot and 
5,920-foot contours shown on the Cold Creek 15-minute 
Quadrangle in the SE '4 Sec. 36, T. 23 N., R. 56 E.; 
in northwestern White Pine County, Nevada (figs. 3, 



8). Clear water; soil bottom; moderate current; little 
vegetation; "not quite lukewarm." Hubbs family, Sep- 
tember 11, 1934 (M34-208); UMMZ 132187 (10, 19- 
31 mm.); 6-foot woven-mesh seine. 

The sample of young fish at Location G9 was 
quickly taken, with the idea of merely ascertaining 
whether the fish here are the same as those in 
the colder springs on the west side of the valley 
(Locations G7 and G8), which were assumed to 
represent an undescribed form. The fish were 
caught in a sheet-piled ditch about 0.6 m. wide. 
The rancher, Billy Moore, as was widely known, 
kept goldfish (Carassiiis ouiatus) of varied hues, 
carp (Cypriiuis caipio). and catfish {Ictaliinis 
sp. ). From afar it had been recommended that 
we examine the spring-fed waters of this ranch. 

We think that we may have sampled the only 
then e.xtant stocks of native fish on the east side 
of the valley and the two populations, perhaps 
all that existed, in the northern part of the west 
side ( the southern part apparently has no waters 
capable of having retained fish life). The two 
western Locations, G7 and G8, probably lie at or 
near the southern and northern limits of the fish- 
inhabited springs. Roughly midway between, at 
Strawberry Ranch (in SW V4 Sec. \0. T. 21 N.. 
R. 55 E. ), we were informed on September i 1, 
1934, by the lady rancher, that her springs, then 
about dry in the extreme drought of that year, 
had contained minnows, but that they had been 
destroyed by freezing a few years previously, 
dying in such numbers that they emitted a terrible 
stench when the ice thawed. Thereafter, she 
said, no minnows were observed. Numerous 
springs between Strawberry Ranch and the spring 
that yielded sample G7 are shown on the Eureka 
15-minute Quadrangle, and we suspect that they 
may prove to contain minnows, probably much 
like those taken at Location G7. 

A streamlet about 1.5 m. wide, with Nustitr- 
liuiu. named Cold Creek on Cold Creek Ranch 
15-minute Ouadrangie, was found to be fishless 
where examined near Simonsen (now Cold Creek) 
Ranch on September 11, 1934. This stream is 
shown on the Diamond Springs 15-minute Quad- 
rangle as arising in Cold Spring on the slope of 



158 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




FiGUKt 30. (;'/7i/ hUoliii iiIhsu. Ironi Hiiiiiholdl Riser s>btcm, and a well differentiated race Ironi IJianiond Valley, 
Nevada. A. Hiimholdt River, near Lovelock (Location G2 ) ; UMMZ 12487.3, no. II: considered topotvpic; female, 
lOX mm. B. Hishop Creek (G4): UMMZ I4l.^2.i, no. 24: male, 7h..^ mm. C. Same field collection: no. 3: female, 
114 mm. D. Sulphur Spring. Di:imond V.illey (ddl: UMMZ I24^>27, no. 31: female. S2..> mm. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 159 



Figure 31. Gila hicolor: two subspecies from Newark Valley. Nevada. A. G. h. newaikensis, near Diamond 
Peak (Location G7); UMMZ 188893, no. 10; holotype, male, 68.0 mm. B. G. h. iicwarkensis. same field collection; 
UMMZ 132185, no. 8; paratype, female, 82.2 mm. C. G. h. citchihi. Fish Creek Springs (GIO); UMMZ 124939. 
no. 4; paratype, male, 114 mm. D. G. h. eucliihi, same field collection; UMMZ 124938; holotype, female, 141 mm. 



160 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Diamond Mountains, near the east base. It is 
mapped as being diverted into an irrigation 
ditch. The apparent lack of native fish in this 
httle stream, as well as in the few other spring-fed 
rills that reach the floor of Newark Valley, is 
probably attributable to the flash floods that oc- 
casionally denude canyon streams in the land of 
rare but torrential waterflow. 

It .seems obvious that this subspecies was de- 
rived from Gila hicolor obesa of the Humboldt 
River system, or its immediate ancestor. Its 
distinctiveness is concordant with the evidence 
that the isolation dates from an earlier than the 
latest pluvial time (pp. 22-25). 

Description and Comparisons. 

Holotype. UMMZ IS8893. a nuptial male 68.0 
mm. in standard length (fig. 31 A). Paratypes, 
UMMZ 132185. all other known specimens (226. 
25-97 mm. long) from same Location (G7. data 
given above), including the adult female, 82.2 
mm. long, that is illustrated (fig. 3 IB). 

This subspecies is compared primarily with 
typical Gila Ivcolor obesa. as represented by pop- 
ulations of the Humboldt River system, and with 
the aberrant races, also referred to G. b. obesa. 
that inhabit Diamond Valley. Those drainage 
basins lie adjacent to Newark Valley. Gila b. 
ncwarkciisis is compared with C. b. ciicbila of the 
Fish Creek Valley division of the basin of pluvial 
Lake Newark in the account of that subspecies. 
The other isolated form treated as a distinct sub- 
species. G. b. isolata of the drainage basin of 
pluvial Lake Clover, is compared later with G. b. 
?u'\\(ukci!sis. 

Sizii. Gila bicolor newarkensi.i is of medium 
to rather small size. The largest specimen in each 
of the two main collections measures 97 (G7) 
and 67 ( GS ) mm. 

Coi oRAnoN. The general color tone is darker 
and more uniform over the body than in the pop- 
ulations referred to G. b. obesa, not closely ap- 
proaching the bicolored pattern of that subspecies. 
The coloration seems to enhance the turgid body 



form. Characteristically, the dark pigmentation 
of the sides is less uniform than in the other 
forms, because the melanophorcs are thickly and 
broadly concentrated around the margins of the 
scale pockets, leaving the rounded central area 
of the pockets largely clear, usually to form rather 
conspicuous stripes along the horizontal scale 
rows (figs. 31A,B). This pattern is usually most 
conspicuous ventrally. In some forms referred to 
G. b. obesa. sparser black pigment tends to mar- 
gin the scale pockets (p. 151), less conspicuously, 
in a thinner, more diamond-shaped pattern, with- 
out forming definite horizontal streaks. The dark 
pigment extends farther down the sides than in 
even the upstream populations of G. b. obesa 
(fig. 30B,C), usually more or less completely 
rounding the caudal peduncle ventrally and reach- 
ing to a narrow midventral light wedge on the 
belly. The underlying dark lateral band is evident 
in small specimens, but seems to fade at a smaller 
size than it does in G. b. obesa. The basicaudal 
spot is replaced by a thin blackish streak along 
the curving posterior border of the squamation 
( in some young, the spot is weakly evident ) . This 
blackish streak may be confined to a vertical sub- 
median position, or may extend farther, curving 
aroLind the dorsal and ventral lobes of the .squa- 
maticMi. As noted above (p. 152). the basicaudal 
spot tends to become minute or diffu.se upstream 
in G. b. obesa. but not to form a streak. The 
fins are all darkened. 

Lirr COLORS. The fish at Location G7 were 
described as follows, and those at G8 were noted 
as having the same color. Relatively uniform 
olive-green above to silvery below. Some have a 
very strong wash of pinkish-brassy, a yellow 
pectoral axil, and coppery-red-brown lower fins 
(whitish in others). 

Form. The head and body arc strongly turgid, 
rounded in all aspects. The contrast with fish 
from Sulphur Spring in Diamond Valley is 
sharpest, because the muzzle is much more 
broadly rounded, in both dorsal and lateral view, 
and the mouth is usually lower, more curved, and 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



161 



O 
O 

q 

X 

_l 250 

CD 


• 

• • • 

• 
•••••• 


• 

• 
+ 


+ + 


+ 

+ 
+ 


+ 

1 


• G b newarkensis 
o G b isolafa 
+ G. b obesa 

+ 
+ + 

1 1 


CL 

X 

^ 200 

z: 

_l 


O O ^H- 

+ 

+ 

1 1 


o + 


+ 
o 

1 


+ 
+ 

+ 



40 



50 



60 



70 



80 



90 



100 



10 



STANDARD LENGTH, mm 



Figure 32. Usual distinction of Gila hicalor newarkensis from G. b. isolata and G. b. obesa in length of pectoral 
fin of males. The G. h. obesa material came from Bishop Creek (Ci4) and from HimTholdt River near Lovelock 
(G2) andCarlin (G3). 



less oblique, becoming more nearly horizontal 
forward; the mandible is slightly included at its 
front tip. The nuchal region is more humped, 
and the dorsal contour is scarcely elevated at 
front of dorsal fin. On the average these distinc- 
tions also hold when G. b. newarkensis is com- 
pared with the other populations referred to G. b. 
obesa. The fins are rounded, without any falca- 
tion, thus contrasting somewhat with even the up- 
stream and Diamond Valley populations referred 
to G. b. obesa. 

Lateral-line system. In half-grown fish the 
lateral line is sometimes more or less interrupted 
posteriorly, but in adults is nearly or quite com- 
plete. 

In both main series of G. b. newarkensis. and 
in G. h. eitchila, unlike all other populations of 
the species studied for this report, the supratem- 
poral canal (table 26) is definitely and charac- 
teristically more often complete than incomplete 
— apparent evidence of the common origin of all 
the Newark populations. 

Morphometry (table 27). On the average, 
and generally with limited overlap, G. b. new- 
arkensis (as also G. b. eiicliila) differs from the 



forms referred to G. b. obesa as follows: the pre- 
dorsal length is greater (confirming the trend 
toward greater development of anterior parts in 
isolated spring forms ) ; the distance between 
pelvic insertion and anal origin is shorter; and the 
body, with .some exceptions, the caudal peduncle 
(especially at Moores Ranch ), and the head aver- 
age deeper. The mouth, as indicated by the 
mandible length, is similar in size to that of G. b. 
obesa. and averages smaller than that of either 
G. />. eiieliita or G. b. isolata (table 27. figs. 34, 
37). 

The lengths of the pectoral and pelvic fins are 
about as usual in females of G. b. newarkensis and 
G. h. enchila, but in the males these fins, es- 
pecially the pectoral, are unusually large (table 
27, fig. 32), larger than in the other populations 
except that from Sulphur Spring. 

Sexual dimorphism and nuptial charac- 
ters. Apparently to a greater extent than is usual 
the dorsal fin is more posterior in females than in 
males, and in this subspecies (and also G. b. 
eiieliila and the Sulphur Spring population of G. 
b. obesa ) the fins, especially the pectoral, are un- 
usually large in males but are of about usual size 



162 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 27. Hropdrlicnal ))U'(isiiiei)H-nls. in pcniiillage of staudanl lcni;lh. far representative scries of Gila bicolor 
in certain basins in Nevada. For eacli entry tliere is given the range, below this llie mean, and, for the standard length 
and for items based on fewer specimens, the number of specimens (as a subscript). (Table continued cm next three 
pages.) 



Subspecies 
Pluvial lake system 

Locality 



Gila bicolor obesa 



newarkensis 



euchila isoluta 



Lahontan 



Diamond 



Newark 



Newark Clover 



Standard length 
Smaller 9 9 

Larger 9 9 

Smaller i i 

Larger $ $ 

Predorsal length 
Smaller 9 9 

Larger 9 9 

Smaller c5 i 

Larger i S 

Anal to caudal 
Smaller 9 9 

Larger 9 9 

Smaller S S 

Larger 6 S 

Pelvic to anal 
Smaller 9 9 

Larger 9 9 

Smaller i S 

Larger i S 

Body depth 
Smaller 9 9 

Larger 9 9 

Smaller c! 6 

Larger S i 



Humboldt Humboldt 

R. nr. R. nr. Bishop 

Lovelock Carlin Creek 



52-94 45-98 45-95 

14,, 6U, 7l,r. 

108-115 100-105 101-137 

112,, 103, Ills 

53-73 48-73 48-76 

63,-. 63,, 61,,. 
79-96 74-107 — 
o3e 82i2 — 



521-562 534-567 529-580 

539 549 558 

548_567 539-541 544-590 

558 540 569 

516-545 525-557 525-569 

533 542 547 

518-540 533-570 — 

527 546 — 



276-322 291-320 289-323 

300 306 309 

285-310 297-305 280-326 

296 301 306 

291-335 292-329 300-341 

314 310 324 

294-328 292-320 — 

309 303 — 



179-208 165-205 174-214 

192 184,,, 145 

177-194 186-196 177-214 

184 191 196 

177-219 166-204 173-202 

191 187 184 

179-204 167-197 — 

189 186 — 



244-299 249-299 269-314 

270 268 287 

263-300 259-269 281-311 

276 264 299 

246-274 249-282 261-313 

263 265 284 

265-293 258-287 — 

279 274 — 



Birch Sulphur Near Dia- Moores 
Ranch Spring mond Pk. Ranch 



Fish Indepen- 

Creek dence 

Springs Valley 



55- 
66, 


-81 


53- 
67„ 


■86 


47- 
53, 


-61 


46- 
59,, 


-70 




- 


75- 
80„ 


-87 



532-559 530-570 
551 554 



520-541 522-574 
532 542 

— 515-556 

— 536 



283-326 277-320 
305 295 



306-336 2X7-324 
321 305 

— 288-321 

— 306 



l,S(l-208 173-206 
1 47 190 



166-207 170-207 

185 189 

— 183-194 

— 188 



274-318 275-315 
300 296 



265-304 243-306 

287 274 

— 277-324 

— 302 



47-97 


49-67 


49-98 


51-91 


63 „. 


55,, 


74,.-. 


68,, 








101-143 


— 


— 


— 


I24,,,, 


— 


50-69 


43-54 


45-73 


47-73 


58,, 


49„ 


59,.. 


63„ 








77-114 


— 


— 


— 


91-, 


— 


543-602 


555-610 


565-621 


560-593 


572 


581 


584 


577 








561-613 


— 


— 


— 


590 


— 


545-588 


545-579 


550-596 


560-610 


569 


563 


575 


578 








559-599 


— 


— 


— 


579 


— 


285-333 


289-317 


249-3 1 1 


251-304 


310 


300 


288 


288 








265-3 1 3 


— 


— 


— 


285 


— 


309-345 


300-337 


286-330 


282-313 


323 


316 


307 


302 








290-3 1 5 


— 


— 


— 


301 


— 


155-185 


158-193 


152-187 


161-201 


170 


172 


170 


185 








165-188 


— 


— 


— 


175 


— 


158-185 


155-192 


160-141 


163-216 


172 


161 


171 


184 








163-180 


— 








lf)8 


— 



281-338 280-316 
304 248 



241-324 277-304 
307 288 



244-310 278-323 

283 303 
271-316 — 

242 — 

263-302 2S4-324 

282 304 
267-3 1 I — 

294 — 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



163 



Table 27. continued. 



Subspecies 




Gila hicolor obi 


?sa 




iiewad 


censis 


ciicliila 


isolata 


Pluvial lake system 




Lahontan 




Diamond 


Newark 


Newark 


Clover 


Locality 


Humboldt 

R. nr. 
Lovelock 


Humboldt 
R. nr. 
Carlin 


Bishop 
Creek 


Birch 
Ranch 


Sulphirr 
Spring 


Near Dia- 
mond Pk. 


Moores 
Ranch 


Fish 
Creek 
Springs 


Indepen- 
dence 
Valley 


Peduncle depth 




















Smaller 9 9 


111-133 

123 


107-131 
119 


112-135 
126 


118-133 
125 


117-133 
125 


120-145 
131 


135-150 
142 


120-142 
131 


1 19-149 
135 


Larger 9 9 


114-132 
120 


119-125 
122 


120-132 
127 


z 


— 


— 


— 


1 18-141 
128 


— 


Smaller S S 


120-135 
126 


105-129 
121 


118-135 
130 


117-138 
130 


115-135 
124 


132-148 
138 


134-157 
145 


123-148 
136 


135-153 
142 


Larger $ S 


119-130 

124 


115-132 
123 


— 


— 


121-137 
130 


— 





120-138 
132 


— 


Head length 




















Smaller 9 9 


275-312 
295 


285-314 
298 


282-329 
299 


290-301 
296 


288-328 
315 


277-309 
294 


283-317 
297 


307-348 
323 


295-333 
312 


Larger 9 9 


282-299 
292 


284-298 
291 


271-319 
298 


z 


— 


— 


— 


294-337 
3 16 


— 


Smaller £ S 


276-306 
288 


285-308 
293 


286-315 
299 


281-304 
290 


294-322 
307 


271-306 
296 


272-300 
291 


300-341 
318 


292-319 
308 


Larger c$ S 


282-300 
289 


275-305 
292 


— 


— 


283-317 
302 


— 


— 


308-324 
313 





Head depth 




















Smaller 9 9 


183-199 
191 


185-211 
196 


194-222 
204 


199-207 
203 


200-223 
213 


203-223 
211 


210-225 

217 


203-237 
220 


208-226 
217 


Larger 9 9 


184-199 
192 


187-194 
191 


190-218 
203 


— 


— 


— 


— 


2 12-232 
224 


— 


Smaller £ S 


178-196 
189 


140-202 
196 


IS5-222 
205 


192-211 
2(10 


191-213 
204 


201-236 
217 


208-228 
217 


213-234 
223 


207-228 
218 


Larger i S 


186-197 
191 


185-203 
193 


— 


— 


188-214 
202 


— 


— 


212-237 
226 


— 


Head width 




















Smaller 9 9 


139-163 
149 


138-163 
148 


150-175 
161 


153-171 
158 


157-191 
170 


151-163 

157 


151-167 
159 


155-195 

17! 


152-180 
168 


Larger 9 9 


146-160 
153 


146-151 
149 


153-181 
168 


— 


— 


z 


— 


167-194 
182 


z 


Smaller S S 


132-149 

142 


139-157 
147 


146-168 

157 


137-165 
154 


149-176 
159 


146-171 
158 


150-167 
159 


156-I7S 
166 


156-186 
169 


Larger S S 


137-150 
145 


133-161 
148 


. — 


— 


149-168 
159 


— 


— 


159-195 
174 


— 


Snout length 




















Smaller 9 9 


68-85 
79 


72-89 
78 


74-91 
83 


79-86 
82 


79-1110 
84 


75-86 
80 


79-101 
87 


76-103 
92 


77-96 
88 


Larger 9 9 


78-85 
81 


76-76 
76 


77-97 
87 


— 


— 


— 


z 


,S6-103 
95 


z 


Smaller S S 


69-85 

77 


71-83 
76 


77-89 
85 


74-87 
81 


75-87 
79 


74-83 
78 


77-87 
83 


84-102 
92 


77-90 
86 


Larger <5 S 


74-82 
78 


71-85 
78 


— 


— 


73-84 
79 


— 


— 


86-97 
92 


— 



164 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



TaHLE 27. CONTINUED. 



Subspecies 




Gila hicolor ohesa 




newai 


■keiisis 


eiichild 


isolata 


Pluvial lake system 




Lahontan 




Diamond 


Newark 


Newark 


Clover 


Locality 


Humboldt 

R. nr. 
Lovelock 


Humboldt 
R. nr. 
Carlin 


Bishop 

Creek 


Birch 
Ranch 


Sulphur 
Spring 


Near Dia- 
mond Pk. 


Moores 
Ranch 


Fish 
Creek 
Springs 


Indepen- 
dence 
Valley 


Orbit length 




















Smaller 5 9 


60-85 
70 


59-88 
76 


58-76 
66 


67-75 
71 


59-84 
75 


60-83 
69 


67-78 
73 


55-80 
67 


54-76 
65 


Larger 9 9 


56-58 

57 


58-58 
58 


46-60 
52 


— 





— 


— 


49-59 
53 


— 


Smaller S 5 


(i8-8: 

74 


67-89 

74 


65-76 
71 


67-85 
75 


74-81 
78 


64-80 

72 


70-81 
76 


64-86 

75 


61-74 

67 


Larger S S 


61-68 
65 


58-68 
63 


— 


— 


63-7 1 
67 


— 


— 


51-70 
63 


— 


Upper-jaw length 
Smaller 9 9 


74-91 
81 


65-88 

79 


76-91 

84 


77-87 
SI 


69-95 
81 


69-87 

79 


70-89 
80 


85-103 
93 


84-98 
91 


Larger 9 9 


78-86 
81 


79-81 
80 


75-94 
87 


z 


— 


— 





84-104 
96 


— 


Smaller c? i 


74-85 
78 


71-85 
79 


77-91 
84 


73-86 
80 


64-86 
79 


71-89 

77 


73-89 
81 


73-99 
89 


83-99 
89 


Larger S S 


75-81 

77 


75-88 
81 


— 


— 


73-85 
79 


— 


— 


86-97 
91 


— 


Mandible length 




















Smaller 9 9 


98-120 
108 


98-12(1 
107 


94- 1 1 1 
104 


1(10-112 
107 


103-123 
114 


92-109 
103 


95-112 
104 


107-125 
1 16 


105-120 
114 


Larger 9 9 


104-117 
109 


101-105 
103 


94-1 IS 
108 


— 





— 


— 


107-122 
114 


— 


Smaller 6 S 


99-1 10 
105 


96- 1 1 2 


96-108 
103 


98-110 
103 


100-111 
107 


>I0-1(I6 
100 


94-109 
103 


105-1 19 
113 


105-119 
112 


Larger <^ i 


97-110 
102 


100-111 
106 


— 


— 


100-113 
106 


— 


— 


105-120 
113 


— 


Interorbital width 




















Smaller 9 9 


89-108 
97 


86-106 

97 


94- 1 1 2 
102 


93-102 
96 


93-113 
108 


S9 104 
9h 


96-114 
104 


100 1 12 

105 


91-107 
100 


Larger 9 9 


91-102 

97 


95-96 
95 


95-105 
101 


— 


— 





— 


V6-105 
100 


— 


Smaller S £ 


85-102 
94 


89-1 1 1 
9X 


94-106 
10! 


9 1-105 
99 


88-103 
97 


84-99 
92 


9S-122 
108 


96-116 
106 


94-108 
100 


Larger i $ 


87-96 
91 


81-99 
93 


— 


— 


91-102 
96 


— 


— 


96-106 
101 


— 


Suborbital width 




















Smaller 9 9 


34-43 
38 


32-42 
37 


32-47 
40 


34-43 
38 


34-50 
44 


35-43 

39 


37-50 
42 


43-57 
4S 


37-46 
42 


Larger 9 2 


36-44 
40 


41-41 
41 


40-46 
43 


— 


— 


— 


— 


45-53 
49 


— 


Smaller i 6 


31-40 

35 


33-42 

37 


34-44 

39 


32-47 
38 


31-43 
38 


33-45 

39 


36-52 
42 


44-53 
48 


38-44 
41 


Larger $ <5 


36-39 
38 


37-43 
39 


— 


— 


38-49 
44 


— 


— 


41-56 
47 


— 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



165 



Table 27. continued. 



Subspecies 
Pluvial lake system 

Locality 



Gila bicolor obesa 



newarkensis 



euchila isolata 



Lahontan 



Diamond 



Newark 



Newark Clover 



Humboldt Humboldt 

R. nr. R. nr. Bishop 

Lovelock Carlin Creek 



Birch Sulphur 
Ranch Spring 



Near Dia- Moores 
mond Pk. Ranch 



Fish Indepen- 
Creek dence 

Springs Valley 



Depressed dorsal 
Smaller 9 9 

Larger 9 9 

Smaller S $ 

Larger S $ 



222-258 205-264 202-238 

243 234,. 220 

212-238 206-227 194-227 
228 217 211 

241-283 222-248 210-256 

255 236„ 236 

227-249 219-238 — 

237 228 — 



220-264 235-286 
2467 259,,, 



245-271) 245-279 

257,,, 262 

— 249-269 

— 257 



212-252 210-242 
236 227 



233-271 232-287 
257 251,, 



205-248 198-249 

223,,, 232 

199-220 — 

209 — 

230-273 218-249 

248 237 

2 1 2-243 — 

229 



Caudal length 
Smaller 9 9 

Larger 9 9 

Smaller 3 S 

Larger 5 S 



263-295 244-271 222-287 

280; 261, 251 

253-264 217 217-243 

2583 217, 2357 

266-317 260-269 234-286 

288, 265; 257 

261-270 258-265 — 

264^ 261,, — 



270-281 241-311 
275. 276,; 



267-273 262-288 
270., 2787 



225-275 249-283 
247,7 265 



241-290 255-300 

258,,, 277,,, 



258-284 



236-273 245-269 

253,. 260,,. 

215-257 — 
239,, — 

248-292 251-278 

261,,, 265,, 

260-265 — 
263, — 



Pectoral length 












Smaller 9 9 


185-221 


173-200 


154-188 


1 84-2 1 2 


197-236 




201 


189,7 


175 


199 


216 


Larger 9 9 


191-206 


169-197 


167-185 


— 


— 




198 


183 


174 


— 


— 


Smaller £ $ 


198-241 


204-237 


186-232 


199-242 


217-252 




223 


215 


209 


225 


234 


Larger 5 S 


217-230 


197-230 


— 


— 


208-253 




223 


209 


— 


— 


234 


Pelvic length 












Smaller 9 9 


158-198 


159-188 


147-165 


169-198 


178-208 




182 


170,,, 


160 


181 


193,,. 


Larger 9 9 


175-183 


157-171 


153-178 


— 


— 




178 


164 


163 


— 


— 


Smaller cS S 


176-208 


171-190 


164-193 


180-21 1 


196-213 




190 


179 


176 


195 


202 


Larger £ $ 


184-201 


164-186 


— 


— 


193-208 




191 


177 


— 


— 


202 



172-204 174-201 
185 189 



238-268 231-287 
250 252 



148-175 145-191 
163 164 



172-199 168-197 
189 183 



167-197 168-200 
187 183 

168-196 — 
183 — 

207-281 179-223 
242 212 

204-235 — 
219 — 



151-179 152-170 
1 64 1 60 

150-170 — 
1 60 — 

164-206 152-176 
184 166 

177-200 — 
187 — 



166 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Table 28. ScmuiI ilnxniplusm in picilorsiil and I'm lcn:^ths in pnpiihiiions of Ciila bicolor /;; ccrtiiin luisins in 
Ncvcitlci, e.xpiL'ssc'cl as excess for males over females in mean values in perniillage of standard lent^lli.^ 



Subspecies 

Pluvial lake system 
Locality 



Predorsal length Dorsal fin 



Caudal fin 



Pectoral fin 



Pelvic fin 



Small Large 
fish fish 



Small 
fish 



Larce Small 
fish fish 



Larye Small 
fish fish 



Large Small Large 
fish fish fish 



Gila bicolor ohesa 

Lake Lahontan 

Near Lovelock (15/23: 6/A)-' 
Near Carlin (12/20: 12/21 
Bishop Creek ( 15/15; —I 

Lake Diamond 

Birch Ranch (11 10: — ) 
Sulphur Spring ( 13 20: — 1 

Ciilti liiioloi luiviirkciisis 

Lake Newark 

Near Diamond Peak ( 1 1/19: —1 
Moores Ranch (14 15; — 1 

Gilii hici'loi ciicliilii 

Lake Newark 

Fish Creek Springs ( 16 25; 5 '13 1 

(Hill I'iiolor isoliild 

lake (lover 
Independence Valley ( 15/15; — ) 



-6 


-31 


12 


-7 


(S 


-\ 


11 


— 


Ifi 


13 


_ 


13 


-8 


— 


3 



S 


6 


->-> 


25 


S 


13 


4 


44' 


26 


26 


y 


13 


6 


— 


34 


— 


16 


— 


-5 


__ 


26 


_ 


14 


_ 


1 


— 


18 


— 


9 


— 



-15 



21 
24 



11 
12 



65 
63 



-33 



24 



5 — 



36 



29 — 



26 
19 



20 



' Based on values in table 27. 

-The ninnbers in parentheses for each Location represent females atid males, respectively, first for the sniuller fish I ni flic size classes of 45-9y 
mm. for females and 4.1-73 mm. for males, mcliuhnii one fish 7h mm, lon^ from Bishop Creel;) and then for the larger adults (luil-14.1 mm. lor 
females, and 74-114 mm. for males). 

■Only one Icmale measureii; other t.dlies for Ihe l.iiuLiI lin .ire (T.isei! on a lediiced number of specimens, because this fin is often broken. 



in fenialcs. The sexual dinioiphiMii in the length 
of the fin.s is e\treme (table 2rS; figs. 31A-D). 
in this lespeet. (J. h. ncwmkcnsls is approxi- 
mately niatehed only by its eongener G. h. 
ciicJuhi. 

The nuptial ehaiaeters of the male holotype 
( fig. 3 1 A ) are eharaeteristic of the whole species 
( p. M'^)) . The tubercles are confined to the upper 
surface of the pectoral fin. where they are 
strongest on the second and third rays, moderately 
strong ak>ng the first ray, and weak on the fourth 
ray. They are tmiserial throughout on the first 
ray and basally on the next three, branching 
once, with the inner branch very weak on the 
fourth ray. They are regularly aligned, one per 
articulation of the ray. Each organ is sharply 
pointed and is slightly hooked niesad. The outer 
ray is rather thickly padded and the following 



several rays are considerably thickened. The fin 
is rigidly expanded in the horizontal plane, with 
the median area arched ujiward. 

Fin RAts (tables 24, 30). The dorsal rays, as 
usual in the Nevada populations of Gila bicolor, 
seldom deviate in number from S. The anal-ray 
count averages low, and is modally reduced to 7, 
111 all the samples from the basin of pluvial Lake 
Newark (further evidence of consanguinity and 
of meristic reduction in isolated springs). The 
caudal rays range from 16-20 at the type Loca- 
tion. The pectoral rays average somewhat less 
at Location Ci7 ( l.^.SS) than at GS ( 16.50) oi 
in G. />. cuchiUi ( l(i.7l I. The pelvic rays are 
more markedly redticcd in average number in all 
the samples from the basin of pluvial Lake 
Newark (means 8.10-8.66) than in the samples 
referred to G. h. ohcsa (9.09-9.49) — again con- 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 167 

Table 29. Nunihcr <</ rays in the vertical fins in populations of Giia bicolor in certain hasins in Nevada. 

Subspecies Dorsal rays Anal rays Caudal rays 

Pluvial lake system 



— 


60 


1 


1 


46 


T 


1 


19 





-) 


50 




— 


63 





3 


61 


6 


7 
1 


62 

8 


1 



1 


23 


— 


25 


18.88 


1 


.5,S 


_ 


59 


18.98 


T 


36 


5 


43 


19.07 


— 


19 


— 


20 


18.90 


1 


23 




24 


18.96 


1 


26 


— 


27 


18.96 


9 


50 


4 


65 


18.85 


S 


29 


3 


37 


1 8.95 


1 


9 


— 


10 


IN. 90 



Locality 7 8 9 10 No. Ave. 6 7 8 9 No. Ave. 16 17 18 19 20 No. Ave. 

Gila bicolor obesa 
Lake Lahontan 

Carson River' — 35 

Humboldt River 
Near Lovelock 
Near Carlin 
Bishop Creek 

Lake Diamond 
Birch Ranch 
Sulphur Spring 

Gila bicolor lu-warkeiisis 
Lake Newark 

Near Diamond Peak 
Moores Ranch 
Warm Springs 

Gila bicolor eiicliila 
Lake Newark 

Fish Creek Springs — 66 3 1 70 8.07 1 52 16 — 69 7.22 — — I 37 — 38 18.97 

Gila bicolor isolala 
Lake Clover 

Independence Valley 6 79 — — 85 7.93 1 118 25 — 144 7.17 -- I H 64 — 73 18.86 

1 Some introgression from Gita bicolor pecunijer. 



Table 30. Number of rays in the paired fins (both sides counted) in populations of Gila bicolor in certain basins 
in Nevada. 

Subspecies Pectoral rays Pelvic rays 

Pluvial lake system 



35 


8.00 


— 7 


27 


1 35 


7.83 


61 


8.02 


— 13 


47 


1 61 


7.80 


49 


8.02 


1 20 


28 


— 49 


7.55 


20 


7.95 


1 12 


36 


— 49 


7.71 


52 


7.96 


1 10 


40 


— 51 


7.76 


63 


8.00 


— 3 


59 


— 62 


7.95 


70 


8.04 


1 63 


6 


— 70 


7.07 


69 


7.90 


1 65 


3 


— 69 


7.03 


10 


8.00 


— 8 


-> 


— 10 


7.20 


70 


8.07 


1 52 


16 


— 69 


7.22 


85 


7.93 


1 118 


25 


— 144 


7.17 



Locality 13 14 15 16 17 18 19 No. Ave. 6 7 8 9 10 11 No. Ave. 

Gila bicolor obesa 
Lake Lahontan 

Carson River' — 1 15 32 14 2 — 64 16.02 

Humboldt River 

Near Lovelock — 

Near Carlin — 

Bishop Creek - 

Lake Diamond 

Birch Ranch — 

Sulphur Spring — 

Gilti bicolor newarkensis 
Lake Newark 

Near Diamond Peak 

Moores Ranch - 

Warm Springs — — — — — — — — — — — 

Gila bicolor cuchila 
Lake Newark 

Fish Creek Springs — — 3 30 30 10 2 75 16.71 — 1 

Gila bicolor isolatu 
Lake Clover 

Independence Valley 1 — 10 4!) 41 12 — 1(14 16.50 — 3 

' Some introgression from Gila bicolor pectinijer. 



- 


13 


f>3 


38 


9 


34 


3ft 


3 


7 


Ih 


15 


-> 


3 


13 


32 


4 


■> 


26 


21 


3 


8 


25 


55 


") 1 


_ 


5 


27 


32 



17 


16.27 


84 


15.46 


40 


15.30 


52 


15.7! 


52 


15.48 


13 


15.S8 


66 


1(1.50 



— 


55 


15 


— 70 


9.21 


_ 


71 


67 


— 138 


9.49 


— 


68 


28 


— 96 


9.29 


7 


259 


1 17 


1 384 


9.24 


_ 


72 


32 


— 104 


9.31 


— 


1 14 


9 


1 124 


9.09 


55 


80 


-) 


— 137 


8.61 


105 


31 


— 


— 138 


8.20 


18 


") 


— 


— 20 


8.10 


44 


9J 


1 


— 136 


8.66 


313 


76 




— 392 


8.19 



168 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



finning both consanguinity and the trend toward 
reduction in isolated spring habitats. 

VERTiiBRAE. The vertebral counts ( table 3 1 ) 
average fewer in the populations from the Lake 
Newark system (38.39 to 39.07) than in those 
referred to G. h. ohcsa (39.14 to 39.86). 

Scale rows. Almost all of the 1 1 scale-row 
counts also average lower in the populations from 
the Lake Newark system than in those referred 
to G. h. ohcsii (table 32). This is particularly 
true of the scale counts around body and around 
peduncle. The means are 44.4-46.7 v.v. 52.2- 
56.5 and 24.6-26.3 vs. 29.6-32.0. respectively. 
These data, as do those for vertebrae, confirm 
consanguinity and reduction in springs. 

GiLL-RAKERs. The rakers in G. h. neu-arkensis 
are outstandingly few (table 22). short (table 
25; fig. 28C), soft, and swollen. The mean num- 
ber is fewer than in G. h. ohesa. though in this 
respect G. h. iwwarkcnsis is closely approached 
by the headwater population of Bishop Creek. 
More strikingly in form than in number, the 
rakers of G. b. ncwaikensis, at both Locations 
where a good collection was .secured, contrast 
with those of G. b. obc.sn. as represented by ap- 
proximate topotypcs from the Humboldt River 
near Lovelock (Location G2). They are opaque 
in preservative and are so fleshy and thick that 
they generally are in contact at the base, despite 
the low number. In G. b. obcwa. in strong con- 
trast, the rakers arc slender, translucent, bony, 
and generally separated at the base. 

There is little overlap in gill-raker length be- 
tween G. b. lU'Uiirkcn.sis (or G. h. vuchihi) and 
the populations from Diamond Valley that wc 
refer to G. b. obesa. The raker-length measure- 
ments tor typical G. b. obesa are definitely inter- 
mediate between the values for G. b. ncwarkcnsis 
and the Sulphur Spring population (table 25. fig. 
28). 

Pharyngeal teeth. The teeth (table 23 ) are 
usually 5 — 4. occasionally 4 — 4. rarely 5 — 5 or 
4—5. 

Derivation of name. This subspecies was 



named for Newark Valley and for pluvial Lake 

Newark. 



Fish Creek Springs Tui Chub 

Gila bicolor euchila Hubbs and Miller. 
(Figure .MCD. ) 

(/■(/(( hicoliir cmhihi Hlusbs and MiLLER, 1972. p. 103 
(diagnosis) . 

This subspecies is almost surely restricted to 
the waters of Fish Creek Springs, in Fish Creek 
( Little Smoky ) Valley, in the southeastern corner 
of Eureka County, central Nevada. This valley 
is merely the southwestern expansion of Newark 
Valley, the sump of pluvial Lake Newark (pp. 
22-26). 

Location GIO. — Fish Creek SjHinifs in the north- 
western part of Fish Creek ( l^ittle .Smoky) Valley (trib- 
utary in tlood to Newark Valley), on the valley flat 
near its west end. between the 6,020-foot and fi. 040-foot 
contours on the Pinto Summit and Hollevue Peak l.s- 
minute quadrangles; in main ditch about 0.5 km. below 
junction of the two main spring-fed branches (this 
places location near center of -Sec. 8, T. 16 N., R. 53 E., 
about 3.S km. west of the Fish Creek Ranch houses); 
near southwest corner of Eureka County, Nevada (figs. 
3. 8). Clear, somewhat bitter, very slightly sulphurous; 
fine shelly sand. mud. clay, etc.; moderate current; 
moderate growth of bulrushes. Poltiiiuit^clon ( broad- 
and fine-leafed I, C/uira. and Uliiciiltiria: 20 C. (air 
27 ). Hubbs family, August 17. 1938 (M38-134); 
UMMZ 124938-39 (517, 18-149 mm.); 15-foot seine 
with '-1-inch square mesh. 

Location CilOA. — Collection made close by in another 
ditch, to make sure thai small lish seen here were not 
Rhinicliihys. Clear water; soil bottom; slight current; 
tlense I'oldiiioiiclon. cf. P. pcctinatus; 24° C. (air 28' ). 
Hubbs family. August 17, 1938 (M38-135); UMMZ 
124940 (26. 15-37 mm.); 6-foot woven-mesh seine. 

Collection M38-134 was made in a ditch 2 — 1- 
m. wide and to nearly 1.0 m. deep. These fish, in 
conformity with their morphology, were mostly in 
mid-water, with few on the bottom. They were 
locally known as "whitefish," as well as "chubs," 
presumably because the original .settler. Fentster- 
maker. whose grave near the springs is marked 
on the Bellevue Peak 1 5-minute Quadrangle, u.sed 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



169 



the term "Weissfisch" by which name the larger 
cyprinids of Europe are known to Germans. 

Isador Sara, the rancher then operating Fish 
Creek Ranch, supplementing other local informa- 
tion (p. 25). indicated in 1938 that about four 
years previously, he had stocked Fish Creek 
Springs with rainbow trout (Salmo gairdnerii) 
and brook trout (Salvelinus fonliiwlis) that he 
had obtained at Roberts Creek Ranch. These 
grew remarkably fast, he said, and had pink 
flesh, but were not as agreeable to eat as the 
chubs. He thought that the trout had not re- 
produced and were probably no longer present. 
One of the springs, intermediate in the southern 
drainage ditch, was found by us to swarm with 
the chubs, some adult but mostly young and half- 
grown. It formed a spring pool, about 10 m. in 
diameter, which had a very deep center, largely 
choked with UlriciiUiria containing great quan- 
tities of amnicolid snails, seemingly like some 
sampled in the ditch. 

No indications were obtained, by inquiry or 
field reconnaissance, that any other waters in 
Fish Creek (Little Smoky) Valley contain native 
fish. The extent of flow in Fish Creek is dis- 
cussed in an earlier section (pp. 25-26). 

Despite the circumstance that Fish Creek 
Springs, the .sole habitat of this subspecies, lies 
in the same pluvial drainage basin as Newark 
Valley, the home of G. b. iwuarkensi.s, we regard 
the two as sub.specifically separable. Although at 
times of torrential floods Fish Creek actually 
debauches onto the playa of Newark Valley we 
think it is highly improbable that there has 
been any gene interchange for hundreds and 
probably for some thousands of years. 

It seems highly probable that the common an- 
cestor of G. h. euchila and G. h. ncwaikeiisis arose 
from G. h. ohcsa when pluvial Lake Newark 
drained into Huntington Creek, a tributary to 
pluvial Lake Lahontan by way of Humboldt 
River. The distinctiveness of G. b. euchila and 
G. b. newarkeiisis from G. b. obesa is consonant 
with the evidence (pp. 23-25) that the isola- 
tion of the Newark basin and of its chub fauna 
dates from earlier than very late pluvial time. 



Description and Comparisons. 

Holotype. UMMZ 124938, an adult female 
141 mm. in standard length (fig. 3 ID). Para- 
types 124939, all other specimens (516, 18-149 
mm. long) from same collection (M38-134; data 
given above, under Location GIO); including the 
largest male, 1 14 mm. long (fig. 31C). The young 
from Location GlOA were not designated as para- 
types. 

Comparisons in the following description are 
almost entirely with G. b. newarkeiisis. because 
the two subspecies, of almost certain common 
origin, share many significant features, and be- 
cause G. b. newarkeiisis is compared, above, with 
G. b. obesa. 

Size. An outstanding feature of this subspecies 
is its large size. The largest specimen measures 
149 mm. in standard length, and many exceed 
100 mm., whereas the largest example of G. b. 
newarkensis is only 97 mm. long, and the next 
largest .specimen among the many hundreds in all 
the populations of all forms treated in this report 
is a 1 37-mm. example of G. b. obesa from Bishop 
Creek. The distinction in bulk is even more 
striking than in length. 

Coloration. One of the most impressive fea- 
tures that are shared with G. b. newarkensis is 
the color pattern, which is essentially the same in 
general darkness and in pattern, involving rows 
of light-centered scale pockets, the far-ventral ex- 
tension of the dark color, the darkening of the 
lower fins (even more pronouncedly than in G. 
/). newarkensis). and the transformation of the 
basicaudal spot into a blackish lining of the 
curved terminal margin of the scaly area. 

Life colors. The field notes indicate that, 
distinctively, the females are deep moss-green on 
the back, with darker scale borders that tend to 
converge backward. The sides are usually 
strongly mottled or .speckled on individual scales, 
with considerable gilt on the lighter scales. Some 
cheek scales show blue reflections. The lower 
sides are olive, abruptly giving way to the white 
ventral surface. The lower fins are deep-olive. 



170 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



o 
o 
o 



X 

o 



350 



340 



330 



MALES o 
FEMALES • 



•HOLOTYPE 




270r 
1 



MALES 
FEMALES 



50 



60 



70 



120 



130 



140 



150 



80 90 100 no 

STA NDARD LENGTH, mm 
Fir.LiRE 33. Usiuil distinction of Gila hicolur ciichUa and G. h. lu-warkcusis in length of head of both sexes; show- 
ing percentages separable on basis of line maximizing differences (fitted b\ e>e). 



grading to blackish on the rays and to ycliowisii 
on the incinbtancs. and have inoic or less in- 
definite whitish borders. The dorsal and caudal 
fins are very dark olive. There is often some 
orange in the axils of the paired fins. 

Although in general similar in life colors, the 
adult males have much more gilt than the fe- 
males on cheeks, opercles, and sides, and the gill 
on the body is somewhat rosy. Blue reflections 
are rather strong on the lower sides, and the scale 
margins on the ventral area are orange-red. 
especially along the tnidline (this color feature 
is hardly evident on the low males). There is a 
considerable wash of lemon-orange on the dorsal 
and caudal fins. The axils of the paired fins are 



rather bright orange and this color is rather strong 
on the interradial membranes. The anal, pectoral, 
and pelvic rays are deep-olive. These fins are 
pale bluish near the border, especially toward 
the posterior angle. 

The unusually bright colors observed in the 
field, on August 1 7, take on added significance on 
the finding that not a single nuptial male is in- 
cluded in the collection of 343 specimens, and 
that none of the females seem to be gravid. 

Form. The body contours are typically much 
less turgid than in G". /'. newtii-koisis and the head 
is much more pointed in side view, with the tip 
much nearer the horizontal midline of head. The 
much straighter and less decurved anterodorsal 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



171 



o 
o 
o 



X 

X 

I- 



Q 

< 
Q 

< 

I- 



125 



120 



I 15 



I 10 



105 



o o 



o 
o o 



o 
oo 



oo 

o o 

o 

o 

o o o 



o o 
o 



HOLOTYPE 

o 



-each I la ( 59) 
o 
88% r 12% 



QD 
Q 

z 

< 100 

Ll 

o 

o 

-z. 

UJ 

_l 

90- 
t 



..^HOLOTYPE 

o o o 



o 1 12% Al 88% 

o ( ^ 



newarkensis (59) 



50 



60 



130 



140 



50 



70 80 90 100 110 120 

STA NDARD LENGTH, mm 

Figure 34. Usual distinction of Gila bkolor cKchila and G. h. newarkensis in length of mandible of both sexes 
showing percentages separable on basis of line maximizing differences I fitted by eye). 



piofilc: the much larger head; the much larger, 
straighter, and more ohUque mouth, with particu- 
larly mas.sive lips and mandible; and the wide and 
flat suborbital and muzzle, combine to produce a 
very different effect, increasingly so in the large 
fish, but still obvious at like sizes. The impre.ssion 
given by these features, and by observations men- 
tioned above, is that G. b. ciichila is a midwater 
feeder, whereas G. b. newarkensis lives more on 
the bottom. 

Despite its massive size (fig. 3 1 CD), the 
mandible, as in G. b. newarkensis. is slightly in- 
cluded within the front tip of the upper lip. 

The fins are hardly falcate, but less rounded 
than in G. b. newarkensis. 

Lateral-line system. In adults, the lateral 



line on the body is complete or nearly so when 
the scales are undamaged. 

As in G. b. newarkensis. in contrast with all the 
other forms studied for this report, the supratem- 
poral canal is much more often complete than 
incomplete (table 26). 

Morphometry. The trend for isolated spring- 
inhabiting populations to have the dorsal fin set 
farther back than usual is carried in this subspecies 
to an extreme, among all the populations included 
in this study — slightly farther back than in G. b. 
neu-arkensis. but much farther, generally with 
little overlap, than in the other subspecies here 
treated ( table 27 ). The distance from anal origin 
to caudal base averages proportionately shorter 
than in G. b. newarkensis and less than in the 



172 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



other subspecies ( except about the same as in G. 
h. isolata). The distance from pelvic insertion to 
anal origin is also low, as in C. b. iwwarkcnsis. 
Because the anal-to-caudal and pelvic-to-anal 
proportions are both low. it was assumed that the 
distance from pelvic insertion to caudal origin 
might provide a quickly usable character. 
The pelvic-to-caudal dimension, measured with 
dividers on about 30 specimens of each Loca- 
tion, was stepped forward to sec how far it 
reaches. In the several populations the dimension 
reached to any point specified below: 

G. h. ohcsa 

Gl. Carson River near Fallon: from rear part to 

front part of snout. 
G2 and G3. Humboldt River near Lovelock and 

near Carlm: from front edge of pupil almost 

to tip of snout. 
G4. Bishop Creek: from rear ot pupil almost to tip 

of snout. 
G^. Birch Ranch. Diamond Valley: from front part 

of eye nearly to middle of snout, usually to 

front margin of eye. 
G6. Sulphur Spring. Diamond Valley: from rear edge 

of pupil to rear part of snout, usually to front 

part of eye. 

G. h. ncwaikeii.sis 

Ci7 and G8. Near Diamond Peak and at Moorcs 
Ranch. Newark Valley: from rear part of eye 
to slightly in I ront of e\e, usually to about 
middle ol eye. 

G. h. ciichihi 

GIO. Fish Creek Springs, Fish Creek Valley: from 
more than an eye's length behind eye to mid- 
dle, rareh I ront edge of eye, usualh to slightly 
before rear margin of eye. 

G. h. isdlcila 

Gil. Warm Springs, Independence Valley: from rear 
margin to just before front margin ol eye, 
usually to near middle ot eye. 

The proportional measurements ( table 27 ) 
verify the impression that in C /'. eiuhlla. typi- 
cally, the head, snout, upper jaw, mandible, and 
preorbital are distinctively large. In these propor- 
tions it contrasts sharply with G. b. newarkcnsis 
and is rather closely approached only by the 
otherwise very different Sulphur Spring poptila- 



tion (G6. which is referred to G. b obcsu). The 
sharpness of the distinction in length of head and 
length of mandible becomes particularly obvious 
by graphical presentation (figs. 33, 34). When 
the entries for the two subspecies are separated 
by a line seemingly maximizing the differences 
(approximately paralleling the trend of negative 
allometry for the head proportions and the es- 
sential isometry for the mandible proportions), 
the separation for the head proportion is 91 per- 
cent for G. b. cuchihi and 97 percent for G. b. 
newarkcnsis, and lor the mandible 88 percent for 
each subspecies. Since neither form shows signs 
of stunting or abnormalities of growth, it appears 
highly probable that the differences in proportions 
are of basic significance. 

The lengths of the pectoral and pelvic fins are 
about usual in females of C. /'. ciichila and G. b. 
newarkcnsis. but in males the paired fins, espe- 
cially the pectoral, are larger than in the other 
populations, except that frotn Sulphur Spring in 
Diamond Valley. 

SiixUAL DIMORPHISM. .Scxual dimorphism in 
position of the dorsal fin and in length of fins 
(table 28) is extremely high, approximating that 
for G. b. newarkcnsis and far exceeding that for 
any of the other subspecies ( another sign of con- 
sanguinity). As noted above, not a single male 
among the many collected was in nuptial condi- 
tion on August 17. This circumstance, and the 
finding of large numbers of young, indicates 
spring or early summer spawning. 

Fin ravs ( tables 29, 30). Dorsal rays 8 to 10, 
normally 8. Anal rays usually 7, as in some other 
spring-inhabiting subspecies of G/7(; bicolor. in- 
cluding G. /'. isolata and all three populations of 
G. /'. newarkcnsis. Pectoral rays 15-19, usually 
I 6 or I 7, averaging 1 6.7 I ( slightly higher than in 
any other population studied for this report). 
Pelvic rays 7-10. averaging 8.66: lower than in 
any population here referred to G. b. obcsa, 
barely higher than in the type series of G. b. 
novarkensis. modally 9 as in that series, but sig- 
nificantly higher than in the two other populations 
of G. /'. newarkcnsis and in G. b. isolata. 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



173 



Table 31. Nimiher of vertebrae in populations of Gila bicolor in certain hcisins in Nevada. 



Subspecies 

Pluvial lake system 
Locality 



Frequencies' 



37 



38 



39 



40 



41 



42 



No. 



Mean 



Gila bicolor obesa 










Lake Lahontan 










Carson R. 


— 


— 


3 


10 


Humboldt R. 










Near Lovelock 


1 


1 


7 


5 


Near Carlin 


— 


— 


7 


8 


Bishop Creek 


— 


1 


10 


12 


Lake Diamond 










Birch Ranch 


— 


— 


4 


15 


Sulphur Spring 


— 


— 


4 


12 


Gila bicolor neuarkensis 










Lake Newark 










Near Diamond Peak 


2 


7 


6 


1 


Moores Ranch 


— 


3 


7 


4 


Warm Springs 


1 


1 


6 


1 


Gilii bicolor ciicbila 










Lake Newark 










Fish Creek Springs 


1 


7 


4 


-> 



Gila bicolor isolata 
Lake Clover 

Independence Valley 



19 



14 

14 
17 
24 

19 
16 



16 

14 

9 



14 



30 



39.86 



39.14 


39.71 


39.58 


39.79 


39.75 


38.39 


39.07 


38.78 



38.50 



38.23 



^ Including hypurai complex as one vertebra and including the four comprising the Weberian apparatus. 



Vertebrae (table 31). The vertebral aver- 
age, 38.50, is slightly lower than in any of the 
populations here referred to G. b. obesa. within 
the range of variation for the populations of G. b. 
newarkensis, and slightly higher than in G. b. 
isolata. 

Scale rows. The counts of scale rows ( table 
32) in this subspecies and in G. b. newarkensis 
are similar, and average for all II categories 
lower than in any of the populations referred to 
G. b. obesa ( except that the predorsal count is 
the same as in the Diamond Valley populations). 
For most of the categories there is little or no 
overlap; for some, the two forms of the Lake 
Newark drainage basin approximately correspond 
with or approach G. b. isolata: in other categories, 
the counts average lower. 

Gill-rakers. The gill-rakers (table 22, fig. 
29), as in G. b. newarkensis, average somewhat 
fewer than in the populations of Diamond Valley 
referred to G. b. obesa and in the headwater 
populations of that subspecies, and definitely 
lower than in downstream populations of G. b. 



obesa. The range ( 10-14) and the mean ( I 1 .77) 
slightly exceed the values for G. b. isolata. 

In length (tabic 25, fig. 29) and in form the 
rakers in G. b. eiichila (Location 10) are less 
consistently extreme than in G. b. newarkensis 
(Locations 7, 8), but usually are not so long, 
slender, and hard as they generally are in G. b. 
obesa (Locations 1-6). In length the overlap is 
extensive, when the comparison is made on the 
basis of permillages of the standard length, but 
this is in part attributable to the almost invariably 
larger head of G. b. eiichila. In form, the rakers 
in G. b. eiichila tend to be flask-shaped, thick, 
and fleshy basally, but to be more slender and 
less fleshy distally than in G. b. newarkensis. 

Pharyngeal teeth. In all 15 specimens 
checked, the teeth (table 23) number 5 — 4. 

Derivation of name. The subspecies name 
was derived by combining the prefix tv. good or 
well, with xtiAos-. lip, in reference to the con- 
spicuously enlarged mouth and fleshy lips. It is 
treated as adjectival, hence given feminine ter- 
mination. 



174 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 






a 



M C J) 



O i- 

— aj 

03 ^ 



<i3' 



QJ i>^ ^ 






j-i — 



I ^- I . 



rr, ^ r 1 -t 



ri 



7^ 

r\r\ 



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^' r- 


^CO 


— VD 
1^. V-, 


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-T ^> 


— 1/- 



— iO 



7^. 

V-, r- 



\,D r^ r^ r^ ^ r^ 



r- >o cc oo oo oo 






ri ri n rt 



1 ^ 


'i'^, 


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r 1 r 1 


r 1 rt 



00 0^ OG OV 






U 



i 5-5 



ri n ri ri 



r\ I -^ 



r I V-, ^- ■ 



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ri ri — ri 



oc I —> 



'^ \c ^ ^ 



sc r^ ^c ^ 



r I — 

7 -T. 7 =^. 



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I 



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-r -r 



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O^ O O ri 



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E 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



175 



Independence Valley Tui Chub 

Gila bicolor isolata Hubbs and Miller. 
(Figure 35A.B. ) 

Gilo hicolor isolata Hubbs and Miller. 1972. p. 103 
(diagnosis). 

This subspecies is almost certainly confined to 
Warm Springs, the only fish-inhabited water in 
Independence Valley, the eastern arm of the bed 
of ancient Lake Clover, in east-central Elico 
County. Nevada (pp. 29-32). Here, it occurs 
with a companion relict, Rliinichtliys osciihis 
lethoporus (pp. 134-140). Giki does not occur 
in Clover Valley, the western arm of the same 
lake bed, where two populations of R. oscuhis 
comprise another endemic subspecies. R. o. 
oligoporiis. The locality for Gila h. isolata and 
R. o. lethoporus is marked Gil and R12 on 
figure 12. 

Location Gl 1. — Warm Springs of Independence Val- 
ley: described under Location R12 lor Rliinichtliys 
oscidiis lethoporus (p. 134). 

First collection (August 25, 1965). UMMZ 18651S 
and 186906 (285. 25-97 mm.). 

Second collection (April 3, 1966). CAS 24567 (61. 
26-64 mm.), collected by Stephen Harold Berwick 
(aided by William Nisbet and Jeffrey Newman), was 
donated by Dr. Robert J. Behnke to the California 
Academy of Sciences. A lO-foot seine with '4 -inch 
square mesh was used in outlet flowage and main pond. 

The second collector of this subspecies. Mr. 
Berwick, expanded a study of his material into a 
class report, which Dr. Behnke has made avail- 
able. In this report Mr. Berwick treated the pop- 
ulation as an undescribed subspecies and dealt 
with its habitat and with the systematics of the 
tui chubs in general, in correlation with the 
paleohydrography of the Great Basin. 

Gila hicolor isolata abounds in the ramifying 
waters of this spring complex, and seems greatly 
to outnumber Rhinichthys oscaliis lethoporus, 
the associated relict endemic. The chub is more 
midwater in habitat than the dace, and is less 
inclined to take quick refuge in the dense vegeta- 
tion. For some time no dace were recognized 



during the first collection. After a few were 
noticed, vigorous seining in the vegetation was 
required to obtain the series of 101 specimens. 
No dace were included in the second collection. 
Field reconnaissance and repeated inquiries 
brought no indication that the cyprinids were 
introduced into the Warm Springs of Indepen- 
dence Valley, or that any fish occur anywhere 
ekse in that valley. The trenchantly distinctive 
characters of the local form of each of the species 
leave virtually no room for doubt that the popula- 
tions are native. 

Description and Comparisons. 

Holotype, UMMZ 186906, an adult female 
85.8 mm. in standard length (fig. 35B), from 
first collection. Paratypes, all other known speci- 
mens, 25-97 mm., listed above, including, also 
from the first collection, the adult male, 64.6 mm. 
long, that is illustrated (fig. 35A). 

The characters of the two endemic minnows 
and the physiographic evidence (pp. 29-32) 
indicate that both were derived from an ancestor 
that inhabited the adjacent headwaters of the 
Humboldt River, of the pluvial Lake Lahontan 
drainage basin. The connection presumably took 
place in the Pleistocene, but not during very late 
pluvial time. The ancestors were presumably 
Gila hicolor ohcsa. of a type very similar to the 
population sampled in Bishop Creek, and Rhin- 
ichthys osciiliis ruhiistiis ( or very similar pre- 
cursors). 

The two endemic subspecies of Warm Springs 
in Independence Valley agree in being more or 
less dwarfed, and in being among the most 
sharply differentiated of the fishes that were 
derived through the disruption of their now 
endorheic basins from the Lahontan drainage 
system. The evidence is presented in the account 
of Rhinichthys oscuhis lethoporus (pp. 134-140). 

Some osteological features of G. b. isolata are 
used as representative of the genus in the com- 
parative account of the skeletal features of 
Relictus (pp. 182-193). 



176 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



177 



This well marked subspecies is compared with 
G. b. obesa and the two other subspecific isolates 
of Gila bicolor herein described, namely G. b. 
neworkensis and G. b. eiichiUi. both in the drain- 
age basin of pluvial Lake Newark. 

Size. Gila bicolor isolata. like the other iso- 
lated spring forms other than G. b. etichila, is 
somewhat dwarfed. The largest male measures 
73 mm. in standard length; the largest female, 91 
mm. The other similarly dwarfed populations are 
those of Newark Valley (G. b. uewarkcusis) and 
those of Diamond Valley ( aberrant races of G. b. 
obesa ) . 

Coloration. The general tone of the pre- 
served specimens is variably dusky. The under- 
lying axial dusky band is strong in the young and 
typically is somewhat more distinctly retained 
than usual in the adult. Individual blackened 
scales are more conspicuous than in most forms. 
The dark color extends onto the lower sides of 
the body, but a wider band is left unpigmented 
on the lower sides than is usual in G. h. uewarkcu- 
sis and G. b. eiichila. In G. b. isolata, unlike 
those subspecies, the pigment almost never rounds 
the ventral surface of the caudal peduncle. How- 
ever, almost all specimens of G. b. isolata have a 
highly distinctive black speck on the midventral 
line at the very origin of the lower procurrent 
caudal rays — a mark that is almost never de- 
veloped in any of the other subspecies here con- 
sidered. In some specimens, fine lines in a verti- 
cally elongated diamond-shaped pattern appear 
on the lower sides (a feature of the Diamond 
Valley populations referred to G. b. obesa). An 
occasional specimen shows the well aligned 
rounded light scale-pocket centers, such as are 
usually developed in G. b. uewarkcusis and G. b. 
eiichila. The basicaudal black spot, about as in 
Humboldt upstream and Diamond Valley popula- 



tions referred to G. b. obesa, is usually more or 
less evident, though much reduced in size and/or 
intensity. Some specimens bear a trace, gen- 
erally weak, of a fine black line rounding the end 
of the squamation on the caudal base (a usual 
feature of G. b. ncwarkciisis and G. b. eiichila). 
An occasional specimen combines this streak and 
the basicaudal spot. 

Life colors. The life colors were not noted 
in the field, and no bright pigmentation is re- 
called. This is probably a plainly colored form, 
and as such would seemingly contrast with G. b. 
eiichila (pp. 169-170). 

Form. Generally the body is well rounded in 
cross section. The profile is little elevated at nape 
or front of dorsal. The snout is moderately 
pointed ( less so than is usual in G. b. eiichihi ) . 
The anterodorsal profile is less rounded and de- 
curved than it usually is in G. b. uewarkcusis. 
About as in G. b. obesa, contrasting with G. b. 
ncwarkciisis and G. b. euchila, the front tips of 
mandible and upper lip are about even; the 
mandible varies from very slightly included to 
somewhat protruding, rather than being slightly 
to moderately included. In contrast with G. b. 
uewarkcusis, the mouth is usually nearly straight, 
and is sufficiently oblique to rise nearly to the 
lateral midline of the head, to about opposite 
lower edge or middle of pupil, instead of to op- 
posite lower edge of pupil or lower part of eye 
below pupil. The straight, oblique, and large 
mouth suggests midwater feeding. In further con- 
trast, particularly with G. b. newarkensis, the fins 
are rather pointed, though they are more often 
somewhat rounded than slightly falcate (the 
nearest trend toward falcation is shown by the 
anal fin). 

Lateral-line system. The lateral line, even 
in the larger adults, is usually incomplete pos- 



FlGURE 35. Types of Gila bicolor isolata and Rcliclus solirariiis. A. Gila bicolor isolata. Warm Springs, Indepen- 
dence Valley, Nevada (Location Gil); UMMZ 186518, no. 23; paratype, male, 64.6 mm. B. G. b. isolata. same field 
collection; UMMZ 186906; holotype, female, 85.8 mm. C. Relictiis solitarius, Kirkpatrick Ranch, Butte Valley, Ne- 
vada (Collection 7); UMMZ 186904; holotype, nuptial male. 60.3 mm. D. R. solitarius. same field collection; 
UMMZ 141518; paratype, female, 89.8 mm. 



178 



CALIFORNIA ACADEMY' OF SCIENCES 



MEMOIRS 



620 
O 

o 
o 

- 600 



X 



< 



0- 



580h 



5 60 



540 



520 



- 

























- 




















,50 /«'"-•, 


f \ o;j3 — 

















X 








,\li^t93'= 


- 


o 
o 






T 


X 


X 








ot>esa 


X 





t' XO - 


x3^ 


^, x + 


x 

X 




+ 




X 
X 




+ 
+ + 










+ 














+ 




„- — ' ~ 


+ > 


+ 

Ik. 


M + 


X 












X 

+ 




+ 




•If- 


X +* 

+ 
+ 


++ 

+ 
+ 


+ 






+ 


++ 

X 


„,^.BOUDT H ^* ^ 


X 

+X+ 


■Ix X + 


+++ 

+ 


+ + + + + 


+ 


+ 
+ 
+ 
+ 


+ 


+ 




r 7--i83^ 


\9l^ 


X 

+ 


X + 


+ 


+ 
+ 


+ 


+ 










'\3T^^° , 




- 




+ 


+ + 


++ + 












Ve.sHoPCBi^^ 
















1 








1 I 





50 60 70 80 90 100 110 

STANDARD LENGTH, mm 



120 



130 



140 



Figure 36. Usual distinction of Gila hicolcr isolaia from G. h. »hcsa in prcdorsal length, in Bishop Creek (G4) 
and in Humboldt River at Lovelock (G2) and Carlin (G3). combined and separated: showing percentage separable 
on basis of line niaximizint; differences (fitted b\ e\e). 



teriorly. lacking at least on the posterior part of 
the caudal peduncle, usually throughout that 
region, and sometimes farther forward, where it 
may he either lacking or interrupted. This is a 
common feature of western cyprinids that are 
restricted to isolated springs (p. I<S!). but the 
lateral line is not more than incipientl) inter- 
rupted in an_\ of the other subspecies of CJ. hicolor 
here considered. 

Strangely, however, this is the only stibspecies 
among those here treated in which the supra- 
temporal canal is regularly complete (table 26). 
In tills respect. G. b. isoUuci is approached by the 
subspecies of the Newark basin. 

MORPHOMI 1 m . With ver_\ little overlap, the 
dorsal fin is farther back than in any of the other 
forms considered, except G. h. ncnurkcnsis and 
G. h. euchila ( table 27 ). The backward position 
of the dorsal fin is one of the characteristics of 
western cyprinids that inhabit isolated springs. 
When the proportional values for predorsal 
length are plotted against standard length ( fig. 
.i6l. and a line separating the entries for the two 
taxa is drawn approximatelv paralleling tlie allo- 



metric increase in this dimension. 90 percent of 
the specimens in the G. b. isolatu sample are sepa- 
rated from 93 percent of the specimens from the 
Htimboldt sNstem. and the entries for the noncon- 
forming specimens are not far beyond the line of 
separation. This is particularly striking when it is 
noted that the sexes are combined in the graph 
and it is recalled that females tend to have the 
more posterior position of the dorsal fin. It is 
further remarkable that such a high degree of 
distinction is obtained, despite the circumstances 
that the fish from Bishop Creek have on the 
average a more posterior dorsal than those from 
Humboldt River ( of the 10 entries for G. b. obcsa 
that lie just on the G. h. isr^Uita side of the separat- 
nig line. 7 are from BLshop Creek). The Humboldt 
River and Bishop Creek specimens are. respec- 
ti\el\ 97 and S2 percent separable from 90 per- 
cent of those representing G. /'. isohita. 

There are other average distinctions in mor- 
phometry. The distance from anal origin to 
caudal base is shorter on the average than in the 
other subspecies, including G. b. Jicwarkcnsis. but 
is similar in this respect to G. b. ciichihi. Bod\ and 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



179 



O 120- 

o 
o 



m 

Q 

< 



I lOr 



X 
H 

z 

Lij 



100- 



90 



o o 
o 



o 

• '-' r-Sd O 

• ++ • -4- + + ^ OO + -t 

• _ * _•■ +• _+(2+ 

+ + + O ++ + 

• -H- • ++ O- + 

• •• • • • • -H.^ + • + -H- 

♦• • • • o "Ci • + 



o G b isolaia 

' G b newarkensis 

+ G b obesa 



_j_ 



40 



50 



60 



70 



80 
STANDAR D 



90 100 

LENGTH . mr 



no 



120 



130 



140 



Figure 37. Usual distinction of Gila hicolor isolata from G. h. newarkensis and G. h. obesa in length of mandible. 
The G. h. obesa material came from Bishop Creek (G4) and from Humboldt River near Lovelock (G2) and Carlin 
(G3). The horizontal line (fitted by eye) separates 83 percent of the G. />. isolaia entries from 88 percent of the G. 
b. newarkensis entries. 



caudal peduncle average deeper than in most 
forms. The head averages a little smaller than 
in G. b. eiichila and the Sulphur Spring population 
of G. b. obesa, but larger than in the other sets 
studied. The mandible averages about the same in 
length as in the Sulphur Spring population of G. 
b. obesa and in G. b. eiichila, but larger than in 
any of the other forms herein reported upon 
(figure 37 compares the length of the mandible in 
G. b. obesa. G. b. newarkensis. and G. b. isolaia). 
The length of the pectoral fin in males is almost 
always less than in G. b. newarkensis or G. b. 
enchila, about the same as in G. b. obesa. 

SE.XUAL DIMORPHISM AND NUPTIAL CHARAC- 
TERS. Sexual dimorphism (table 28) in the posi- 
tion of the dorsal fin ( indicated by the predorsal 
length) is slight, and the difference between the 
sexes in the size of the dorsal, caudal, and pelvic 
fins is unusually slight. In respect to the size of the 
pectoral fin, the sexual dimorphism averages 
about as in G. b. obesa. and is much less than in 
G. b. newarkensis and in G. b. eiichila. 

The nuptial characters are typical of G;7(; 
bicolor. A small, obviously late maturing male 
46.6 mm. long, in the first collection, of August 
25, exhibits the typical nuptial tuberculation and 



the usual structural modification of the pectoral 
fin. The tubercles are strong on the first 3 rays 
and are arranged one per segment of the ray: 
each is a slender spine hooked basad and mesad. 
They are aligned in one row only on the outermost 
ray and branch once on the second and third 
rays. There are a few on the fourth ray. with 
the file not branching. The tubercles on the 
four rays vary from strongest on the second ray 
through intermediacy on the third and first rays to 
weakest on the fourth. The outer pectoral rays 
are considerably thickened. No tubercles are ap- 
parent on other fins, on the head, or on the body 
scales. Most of the adult males in the second 
collection, presumably yearlings taken closer to 
the normal breeding season (on June 18), are in 
nuptial condition. The tuberculation is similar, 
except that the spinelets extend, decreasing in 
strength, onto additional rays, to the seventh. 
The small size of the nuptial specimens suggests 
that the males first spawn as yearlings. 

Fin ravs (tables 29, 30). Dorsal rays num- 
ber 8, rarely 7. As in G. b. eiichila and in two of 
the three populations of G. b. newarkensis. anal 
rays are predominantly 7, rather than 8. Caudal 
rays deviate downward from 19 in 8 percent of 



180 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



the specimens counted. Pectoral rays number, on 
the average, slightly on the high side. As also in 
two of the three populations of G. h. iicwarkcnsis. 
the pelvic-ray counts are predominantly reduced 
from 9 to 8. 

Vertebrae. The average number of vertebrae 
(table 31) found for C. h. isolatu (38.23) is 
slightly lower than for G. b. newarkcnsis and 
G. h. fiicliild. and definitely lower than in the 
populations referred to G. h. obesa. The modal 
number is 38, instead of 39 or 40. 

Scale rows. In correlation with the reduced 
number of vertebrae, the number of scales in the 
11 rows counted is low (table 32). The mean 
number of scales in the lateral line (48.1) 
roughly corresponds with the means (47.3-49.2) 
found for G. b. ncudikensis and G. b. ciicbila. 
respectively, and is definitely lower than the 
means (52.5-37.3) for the populations referred 
to G. b. obesa. The mean scale count around 
caudal peduncle yields similar differences ( in the 
same sequence. 26.6 compared with 24.6-26.3 
and contrasted with 29.6-32.0 ). The mean num- 
ber of scale rows around the body (51.7) is 
higher than the means for G. b. ?u'\varkciisis 
(44.4 and 45.6) and for G. b. eiidvhi (46.7), 
and is more nearly in agreement with the means 
(51.0-56.5) for G. /'. obesa. Means for other 
row series are intermediate between those for G. 
b. newarkensis and G. b. euchila. as compared 
with G. b. obesa. 

GiLL-RAKERS (table 22). The rakers in G. b. 
isolata average slightly fewer (11.14, with range 
of 8-14) than in G. /). iic\varke)jsis and G. /'. 
eiichila ( 1 1.75-12.23, with range of 9-15), and 
not much lower than in the Bishop Creek head- 
water population of G. b. obesa (12.61, with 
range of 10-15). There is only limited overlap 
on the counts for the Diamond Valley populations 
or on the counts for the other series of G. /'. obe.sa 
considered. 

In length (table 25) and form, the rakers of 
G. b. isolata. except for extreme variants, are 
essentially like those of typical G. b. obesa 
(p. 153), and contrast with the short, fleshy 



rakers of G. b. eiichila and, more strikingly, with 
those of G. b. newarkensis. On the other side, 
the contrast is also great with the long, slender, 
and hard rakers of the populations, referred to 
G. b. obesa, that inhabit Diamond Valley, es- 
pecially with the Sulphur Spring series. 

Pharyngeal TEETtL All 13 specimens checked 
have the expected number of teeth, 5 — 4 (table 
23). 

Derivation of name. The name isolata, of 
obvious significance, was derived from insula -f 
ato. modified in spelling as in Italian, French, 
and English. 

GENUS RELICTUS HUBBS AND MILLER 

Hcliitus Himns and Miiiik, 1472, p. 101 (diagnosis). 

T'l PL SPECIES. Reiicliis solitarius Hubbs and 
Miller. 

Specimens of this cyprinid were first collected 
in 1934 and 1938, when, in the hope of deter- 
mining whether the fish might indicate a Lahon- 
tan or a Bonneville derivation, we first sampled 
the fish life of the endorheic basins of Nevada 
that intervene between those drainage basins. In- 
stead, we found in that area a unique fish in many 
respects unlike any occurring to the west or east 
(or to the north or south). In our treatment of 
the fish life of the Great Basin ( Hubbs and Miller. 
I'^MSb. pp. 51-55). we mentioned this fish "as 
a type of dace that is so distinctive as probably 
to warrant generic separation from Rhiiiiclitliys," 
and, in discussing its limited distribution, we sug- 
gested that the basins it inhabits may be the 
remnant of an ancient, early pluvial drainage 
system. The only subsequent mention of the fish 
in the literature has been a very brief reference 
by La Rivers ( 1962. p. 86) to our 1948 treat- 
ment. 

In reconsidering this distinctive minnow, we 
find considerable support for our long-felt opinion 
that it should be segregated in a distinct genus. 
Until we carried out this renewed study, our 
opinion was based on the indications that it does 



VOL. VII HUBBS. MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



181 



not fit any of the recognized genera, and that it 
displays a distinctive combination of the char- 
acters that had generally been utilized to distin- 
guish genera among western American cyprinids. 
In view of the tenuous nature of the long-used 
characters, which have been falling into some 
disrepute, and in view of the finding of some 
trenchant differences in skeletal structures among 
western minnows (Uyeno, 1960; Uyeno and Mil- 
ler, 1965), we undertook, toward the elucidation 
of this and other problems, a comparative study 
of some bones of the cranium, upper and lower 
jaws, mandibular and hyoid arches, pectoral 
girdle, and fifth branchial arch. The number and 
morphology of the chromosomes has also been 
determined. This study has disclosed a character 
complex of distinctive features, which in our 
opinion confirm, even in this era of lumping, our 
long-held view that the relationships of this 
species are best portrayed by assigning it to a 
separate genus. 

Relictus takes its place, along with Moapa and 
Eremichthys (Hubbs and Miller, 1948a), among 
the distinctive endemic relict fishes that occupy 
limited areas of spring water within the Great 
Basin. It is less extremely restricted in its distribu- 
tion than those genera, for it is native throughout 
most of the basin-bottom springs of four valleys 
(pp. 196-226): Ruby and Butte valleys, within 
the connected pluvial drainages of lakes Frank- 
lin and Gale, and Goshute and Steptoe valleys, 
within the connected pluvial drainages of lakes 
Waring and Steptoe. 

This type of dace provides an excellent example 
of the modification, in part degenerative, that 
characterizes fishes, particularly cyprinids, that 
are confined to isolated springs in the Great 
Basin and elsewhere (Hubbs, 1940, p. 201: 
1941a: 1941b, p. 187: Hubbs and Miller, 1948b, 
pp. 51-52). These modifications include: a 
body form adapted to midwater swimming in 
quiet water, with more or less symmetrically 
curved dorsal and ventral contours of body and 
head; generally rather chubby, with deep caudal 
peduncle: a terminal and generally rather large 



mouth, fitted to engulf organisms living at various 
depths: small, weak, and rounded fins, adequate 
for limited locomotion in quiet water; reduction or 
obsolescence of the barbels, the lateral line, and 
other dermal sense organs, not critically needed 
in the absence of most enemies; scales tending to 
lose their orderly arrangement and wide imbrica- 
tion, and, in correlation, tending to develop radii 
on all fields; (generally) dusky colors. The type 
of dace under treatment (fig. 35C,D) partakes of 
all these modifications. 

Diagnosis 

A cyprinid of moderate size ( larger than 
Rhinichthys), with some distinctive osteological 
characters: dorsal crest of maxilla greatly ex- 
panded upward and backward: cleithrum slender; 
supraethmoid elongate, slender medially but 
notably expanded laterally at front ( resembling 
that of Rhiniclithys): urohyal long and narrow. 
Vertebrae 35-39. Pharyngeal arch moderately 
strong and heavy, but rather thin and somewhat 
lacy on the strongly expanded median section; 
not strongly elevated at the posterior end of the 
tooth row: without a flattened shelf on which a 
second tooth row might develop: teeth 4 — 4 
(rarely 5 — 4 or 4 — 3). Gill-rakers small and 
few (7-12, usually 8-1 I, on first arch). Mouth 
oblique and terminal, completely lacking horny 
cutting edges: no frenum or barbel. Lateral line 
obsolescent, rarely extending to below origin of 
dorsal fin, commonly disrupted: total pores 3-29. 
Supratemporal canal seldom complete ( only 4 of 
76 specimens have the commissure closed), with 
usually 3 or 4 (0-5) pores in each lateral seg- 
ment: prcoperculomandibular pores 11-19; man- 
dibular pores 3-8. Scales rather small (50-70 
transverse rows), poorly imbricated and markedly 
irregular; each usually vertically oval, but some- 
times becoming rectangular with age; with numer- 
ous radii on all fields (much as in Rhiniclithys 
and some other western genera). Fins small and 
strongly rounded; the pelvic especially and 
uniquely paddlelike; dorsal and pelvic both dis- 



18: 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



placed backward, and both beginning at approxi- 
mately the same vertical (as in the subgenus 
Sipluitck's of the genus Giki and in many species 
of the typical subgenus Gilii ) ; dorsal and pelvic 
rays typically 8, anal 7. Nuptial tubercles form 
a highly distinctive pattern (p. 221) on head: 
the largest uniserially line the infraorbital sensory 
canal and suborbital margin; large uniserial 
caducous cones (much stronger than in Gila) 
line the upper edge of the first pectoral ray, 
smaller cones, also strictly uniserial (not forking 
once as they do in Rliiiiichtliys) occur along one to 
several following rays; in high males some 
tubercles develop along outer pelvic rays and 
along first anal rays. Head and body turgid. 
Coloration much as in Siplniteles. rather even, 
and often with large melanophores on lower side; 
lacking the two lateral bands, the head stripe, the 
paired light spots at caudal base, and other fea- 
tures characteristic of Rliiiiichtliys (pp. 104- 
lO.'^). Intestine forming a single, simple, com- 
pressed-S loop (type I of Kafuku. 1958, p. 56). 
as in Rliiniclilliys and many other American 
cyprinids. 

Description 

In this section we expand on the Diagnosis, 
particularly on the osteological characters, while 
largely avoiding repetition. Some additional de- 
scription is presented in the species accoimt. in 
the osteological treatment, we have stressed com- 
parison with Rliiiiichtliys. utilizing primarily 
skeletal material of Rhiiiichthys osciiliis leliqiiiis. 
but with characters to a large degree confirmed 
by examination of skeletons of additional sub- 
species of R. osiiiliis and of other species referred 
to the genus {R. ciiinructac. R. julcatiis. and R. 
titratulus) . 

Skeletal characters, in the oro-mandibu- 
lar region, the maxilla (fig. .^8) proves to be one 
of the distinctive bones of Relictus. This paired 
dermal structure lies dorsolateral to the premaxilla 
and is partly covered laterally by the ventral part 
of the lachrymal, which overlies the entire dorsal 



crest of the maxilla. At its anterior end the 
maxilla is expanded and. projecting from it 
ventromesially. is the rodlike rostral process 
which, with the lateral plate, holds the anterior 
part just behind the a.scending process of the 
premaxilla. The strongly elevated dorsal crest, 
lying midway in the length of the maxilla, is so 
greatly elongated as to occupy about one-half 
that length. Its shape varies from that shown, in 
which the leading edge slopes backward and the 
posterior border is strongly concave, to a form 
with an abruptly elevated leading edge and with 
the posterior border only slightly concave. In 
the considerable expansion of the dorsal crest 
Relictus is unmatched by any other genus of 
western minnows. 

The premaxilla. also a paired dermal bone, is 
overlapped laterally by the maxilla, but is here 
drawn separately (fig. 38B). The anterodorsal 
part forms an ascending ramus cir rostral process, 
which is directed toward the anterior end of the 
cranium, where it contacts the tiny rostral bone. 
The posteroventral end of the premaxilla overlies 
the slightly expanded posteroventral flange of the 
maxilla. The slender rostral process is tilted 
slightly forward at its dorsal end. The slender 
premaxilla tapers gently posteriorly. Its depth 
across the angle at the anterior end is greater 
than one-fourth the total length of the maxilla. 
Its shape is somewhat intermediate between that 
of Gila hicolor and that oi Rliiiiichtliys osciiliis. 

The olfactory region of the cranium contains a 
bone, the supraethmoid (fig. 39B. SE), which, in 
combination with other features, is distinctive of 
Relictus. though it is very similar to that of 
Rliiiiichtliys. The thin, horizontal, unpaired 
supraethmoid sutures posteriorly with the antero- 
medial parts of the frontals ( F ) . Its anteroventral 
part is fused with the anterodorsal surface of the 
underlying ethmoid bone (E). The noteworthy 
featme of this bone in Relictus is its elongate and 
relatively narrow shape: the least width enters 
the total length about 5-8 times. Among western 
cyprinid genera, the supraethmoid of Relictus 
resembles closely only that of Rhiiiichthys. which 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



183 






B 





1 mm 




Figure 38. Lateral (left) and mesial (right) views of left maxillae and premaxillae of adult females of: A. 
Gila bicolor isolata (paratype. 88 mm., UMMZ 186518): B. Relicius soliiaiiiis (91 mm.. UMMZ 177095); C. Rliin- 
ichrhys osciiliis reliqiius (paratype, S5 mm., UMMZ, 1 249t)7 ) . DC, dorsal crest. 



varies considerably in shape among the several 
species but is also elongate and is very to moder- 
ately narrow (checked in the five currently rec- 
ognized species). As seen in dorsal view, the an- 



terior end of the supracthinoid of Relicius is 
incised by a median notch, of variable width and 
depth, that more or less divides this region into 
two lobes, as described for the genus GiUi and 



:84 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




FiGUKi. 3*^. Anterior end of crania of adult females of: A. GiUi himlar isoUiui (p<irat\pe. !S8 nini.. LIMMZ 
1 865 1 (S); B, Rcliclus sdlilaniis (88 mm., UMMZ I77()"^^5); C, Rliiniclilliys osciihis icliqiiiis I paratype, 85 mm., 
UMMZ 124407). E, ethmoid; F, frontal; LE, lateral ethmoid; PE, preethmoid; SE, supraethmoid; V, vomer. 



its relatives ( Uyciio, 1960). The shaft of the 
stipiaethmoid is slenderest medially, but expands 
ni)tably forward (only slightly or hardly at all 
posteriorly). In all other western cyprinid 
genera, the supraethmoid is shorter and broader 
than it is in either Rhinichthys or Relictiis. 

One bone of the hyoid region, the hyomandibu- 
lar, is illustrated for eomparison of Rcliclus with 



Gila and Rliiniclilliys (fig. 40). This is a well 
developed paired cartilage bone that forms the 
posterior suspensory mechanism for the opercular 
apparatus, mandibular arch, hyoid arch, and 
branchial arches. It lies in front of and articu- 
lates posteriorly with the preopercle and opercle 
and dorsally with the hyoinandibular fossa of the 
cranium, its form often provides features of 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 185 



PDC 





B 






1 mm 

Figure 40. Lateral (above) and mesial (below) views of left hyomandibulars of adult females of: A. Gila 
hicolor isolata (paratype. 88 mm., UMMZ 186518); B, Relktiis soUlarius (88 mm., UMMZ 177095): C, Rhinidilhys 
osciiliis reliquiis (paratype, 85 mm., UMMZ 124907). AC, anterior condyle: AW, anterior wing: LHF, lateral 
hyomandibular foramen: LR. lateral ridge: OC, opercular condyle; PDC, posterodorsal condyle; PF, posterior flange; 
VR, ventral ramus, length. 



taxonomic value at the species, or even the 
generic, level, as has been shown by Uyeno and 
Miller ( 1965, fig. 4), whose nomenclature of the 
parts of the bone is here adopted. In Relictiis, 
the anterior wing is moderately to .strongly de- 



veloped. Its slightly convex anterior border ex- 
tends from less than one-third to about one-half 
the distance down the ventral ramus. The dorsal 
edge, between the posterodorsal condyle ( PDC ) 
and the anterior condyle (AC), forms an angle 



186 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




FiGLiKi: 41 . I,atcr;il views of left lower jaws of adult females of: A, CiUi hicolor isolata (paratype. 88 mm., UMMZ 
186518); B. Rcluliis solUarius (41 mm.. UMMZ 177095); C. Rhinklitlixs oscuhis leliqiius (paratype. 85 mm., 
UMMZ 124'-)()7). AF, anterolateral foramen; AR, articular; C'P, coronoiJ process; D. dentary; RA, retroartieular. 



of about 60 with the vertical, as in Gihi. The 
lateral hyomandibuiar foramen (LHF) is at the 
end of a canal roofed by very thin bone that is 
lost in the specimen drawn, thereby giving the 
erroneous suggestion of a difference between the 
position of the opening in Rclictiis and in other 
genera. The posterior flange (PF), which is 
directed obliquely outward from the ventral 



ramus, is rather weakly developed, leaving all of 
the opercular condyle (OC) exposed in lateral 
view. A line drawn from the apex of this condyle 
to the posteroventral tip of the ventral ramus does 
not intersect the posterior flange. The lateral 
ridge varies from moderately developed to nearly 
obsolete, as in the specimen illu.strated. The area 
between the posterodorsal and opercular condyles 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



187 




Figure 42. Mesial views of same lower jaws shown in figure 41. A. angular; AR. artieular; CM. coronoineckelian 
(sesamoid articular); RA. retroarticular. 



is long and broadly concave, as in the subgenus 
Siphdteles. 

On the mesial side of the hyomandibular. the 
position of the dorsal opening of the lateral 
hyomandibular foramen lies opposite or slightly 
below the level of the occipital condyle, well 
within the upper part of the bone, as in Rliinicli- 
tliys iosciiliis). whereas in subgenus Sipluitclcs 
(Gihi hicolor) this foramen opens close to mid- 



length of the hyomandibular, well below the level 
of the occipital condyle. 

The lower jaw is notably larger in Rel ictus 
than in any species of Rliinichthys ( see figs. 
4 1 B.C. 42B,C), and the mandibular joint is less 
concealed by soft tissue ( the mandible was not 
measured in Rliinichthys because the posterior 
end is less readily perceived than in most genera ). 
The dentarv. which forms the anterior and cen- 



188 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




B 




1 mm 



FiGUKE 43. Lateral (above) and mesial (below) views of Icll cleithra ot adult females ot: A. Gila hicolor isolata 
(paratype, XX mm., UMMZ 186518); B, Relktus solitanus (91 mm.. UMMZ 177095); C, Rhinichthys osculiis 
reliquus (paratype. 85 mm.. UMMZ 124907). AS. anterior shelf; DP, dorsal process; HR. horizontal ramus, length: 
K, keel; LR, lateral ridge; VR, vertical ramus, length. 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



189 



tral parts of each lower jaw, curves mesialiy at its 
anterior end to meet, at the midline, the anterior 
end of the opposite dentary. The bone bears a 
prominent ascending coronoid process (CP), 
which, in Relictiis. lies well back on the dentary: 
the distance between the posterior end of the 
angular (A) and the rear margin of the process 
(at the level of the angular) enters more than 
three times in the length of the lower jaw. The 
coronoid process varies from narrow to broad 
and its anterior edge is slightly to strongly in- 
clined posteriorly (as in Gila copci). Running 
along the ventrolateral surface of the dentary is 
the anterior part of the mandibular sensory canal. 
Lying laterally along the midside of the bone is 
the anterolateral foramen (AF): in Relictiis this 
foramen lies well in advance of the tip of the 
angular, as it does also in Gila but not in 
Rhinicluhys (figs. 4IA-C). 

The angular is long and pointed, as in Gila. 
Its posterodorsal surface is notched to receive the 
articulating surface of the head of the quadrate. 
Directly below the notch is a ventral excavation in 
which the retroarticular bone (RA) lies. The 
coronomeckelian bone ( CM ). or sesamoid articu- 
lar, lies on the medial face of the angular near its 
midpoint and slightly below its dorsal margin. 

The cleithrum (fig. 43B), a dermal bone, is 
the largest of the elements that comprise each 
pectoral girdle. In lateral view, the bone ap- 
proaches the form of a reversed L, with the verti- 
cal ramus (VR) longer than the horizontal 
ramus (HR). The cleithrum bears a posterior 
winglike expansion, which is broadest near the 
posteroventral corner of the vertical ramus. A 
shelflike anterior expansion ( AS ) is developed 
from the horizontal ramus, which bears a keel ( K, 
labelled "dorsal edge" by Uyeno and Miller. 
1963, fig. 2) near the mesial edge. A narrow 
space mesial to this keel is partly sutured with 
the coracoid. Mesialiy, the anterior part of the 
cleithrum borders an oval interosseous space and 
contacts the coracoid at its tip. The lateral ridge 
(LR) of the cleithrum, which originates at or 
near the dorsal process ( DP), is slightly deflected 



over the groove that lies along the anterodorsal 
part of the vertical ramus. Both the anterior shelf 
and the posterior expansion are relatively narrow 
in Relictiis. and the anterodorsal flange is short 
and notably expanded. In general, the cleithrum 
of Relictiis is rather slender, because the lateral 
shelf and anterodorsal flange are only moderately 
broadened. 

The median urohyal (fig. 44B) is divided an- 
teriorly into rather short left and right branches, 
each of which is usually further differentiated into 
dorsal and ventral divisions. A slender neck pos- 
terior to this bifurcation is immediately followed 
by three thin, backward-expanded wings, one 
vertical and two horizontal. The laterally ex- 
panded horizontal wings form the ventral surface, 
which is narrow-elongate, and concave along the 
midline. The depth of incision of the bifurcation 
varies from that drawn to at least twice as deep, 
and the width of the ventral surface may be a 
little greater than illustrated, but not nearly as 
broad as shown for Gila bicolor (fig. 44A). The 
urohyal is quite unlike that of Rhiiuchthys (fig. 
44C). 

Pharyngeal arch and teeth. We follow 
the nomenclature of Uyeno ( 1961, pp. 332-333, 
fig. I ). Each lower pharyngeal arch (fig. 45B) 
is strong and moderately heavy, about as in Gila. 
definitely more massive than in Rliinichthys. 
Neither limb is markedly flattened; the anterior 
limb (AL) is shorter than the posterior limb 
(PL). The anterior angle (AA) on the convex 
edge is usually sharply produced and the surface 
toward the angle is notably flat, with a very thin 
edge. The characteristically deeply pitted surface 
( P), through which blood vessels and nerves pass, 
is so very thin that in some individuals either 
one or both arches may be perforated by a small 
to large foramen, or by several foramina. The 
rather swollen convex face near the base of the 
teeth lacks any trace of an alveolus and docs not 
provide a flat enough shelf on which a tooth could 
develop. A posterior angle ( PA ) also tends to be 
developed. The anterior edentulous process is 
shorter than the posterior edentulous process, or 



190 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 






B 





1 mm 




Fir.iiKH 44. Ventral and lateral vievss ol the iirohyaK ol adult females ot : A. Gilii hicolor isnlaia (paratype, 74 
mm., UMMZ ISd.'^I.S); B. Rvhcius soliuiiiiis (Xl mm., UMMZ 177095); C, Rhinuliiln\ ,>st iihis sLihspecies (75 mm., 
UMMZ 17.^790. from l.ittle Bear Fiiver. Utah), 



equally king. Tooth I rises relatively elose to the 
front edge of the posterior limb. The dentigeious 
part of the areh differs sharply from that of sub- 
genus Siplmlch's in not being markedly elevateel 
posteriorly. 

The pharyngeal teeth are all heavy and are 
moderately to very weakly hooked, or the an- 
teriormost one ( IV, represented by its alveolus in 
speeimen figured) may lack a hook in large in- 
dividuals. There is a definite grinding surface on 
most of the teeth. The tooth formula (table 33) 
is normally 4 — f, with occasional noteworthy 
variation ( p, 93 ) to 4 — 3 and 5 — 4 ( with the 
higher number, as usual, on the left side). When 



Taiu.E 3.-!. Ilciihil jiirnuilds in tciliim Cullcclions of 
Relietiis solitariiis Irani ilijjcrcnl dnnniV^c basins. 



Pluvial lake system (Coll. no,)- 
Valley 



Formula' 



0,4—3.0 0,4—4,0 0.5—4,0 



Lake Iranklin 

Ruby ( 1-3, .S, 6) — 9 

Butte ( 10. 1 I ) 1 II 

lake tiale ( 13 ) — 5 

lake Wariug ( 16, 17. 14) i 14 

lake Steptoe (22, 2?, 27, 2S) — 15 

lake Spring (33, 34) — 17 

TOTAL 2 71 



' Counted left-riylit; alveoli included. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



191 




Figure 45. Dorsolateral (left) and mesial (right) views of pharyngeal arches and teeth (or alveoli) of adult fe- 
males of: A. Gilii hicdiiir isolata (paratype, .SS mm.. UMMZ ISfi5l8); B. Relkitis solitarius (91 mm., UMMZ 
171095): C. Rhinichthys ii.sciiliis leliqiiiis (paratype, 85 mm.. UMMZ 124^07). AA. anterior angle: AL. anterior 
limb; P. pitted surface; PA. posterior angle; PL, posterior limb; i-lV, teeth of major row in the arch of Relicins 
(IV represented by alveolus); in the arch of Gila, the first tooth, and in the arch of Rhinichrliys. the first and fourth 
teeth of the major row. arc represented by alveoli. 



192 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 




7 X.*Sy VvV i»5SS re c . -• ^* 9k 



B 



« "vS *J^ '"' 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



193 



only three teeth are developed, the arch ascends 
abruptly to the base of both tooth I and tooth III, 
leaving no flattened space for another tooth, and 
showing no sign of an injury. 

Scale structure. The scales (fig. 46B), 
typically, are vertically oval, with the focus far 
removed from the posterior edge ( nearer the front 
edge). The circuli are rather regular and persist 
along the anterior field, in which, due to the small 
size of this field, they are notably crowded. Radii 
are numerous on all fields. The development of 
basal radii is correlated, as usual, with the rather 
deep embedding of the scales. Among western 
cyprinids, radii are developed on all fields of the 
scales in all species of Rhinicluhys ( though weak 
or obsolete on the basal field in the very distinctive 
R. jalcatiis) and in the monotypic genera (as cur- 
rently recognized) Acrocheilus. Agosia, Eremich- 
thys, Moapa, Orthodon, and Tiaioga; also in 
some species of Giki (in both subgenera, Gila 
and Siphatek's) and, to a varying degree, in some 
forms of Hesperoleucus. 

PsEUDOBRANCHiAE. The pscudobranchiac 
though rather long are usually more or less com- 
pletely bound down to the opercular tissues. This 
same arrangement we find in Gila bicolor and 
Rliinichthy.s o.sciiliis. and also in Moapa coriacca 
and Eicmicluliys acres, contrary to our statements 
(1948a, pp. 2. 15) that the relict genera Moapa 
and Eremichthys lack pseudobranchiae. 

Relationships 

Rclicliis. as indicated previously, has several 
distinctive osteological traits, of which at least 
one (the narrow supraethmoid; fig. 39) is most 
similar to that of Rliinichthys (table 34). The 
dental formula could have been derived from that 
of Rhinicluhys or, just as plausibly, from Gila. 
The lack of a frenum and a barbel, and the 
presence of radii on all fields of the scale, have 



limited value in determining relationships be- 
cause these structures may be lost or indepen- 
dently derived among surviving populations of 
several genera of western cyprinids. The moderate 
vertebral number may simply represent the reten- 
tion of a generalized condition. The pigmentation 
(described on pp. 209-212), bears a striking 
resemblance to that of the subgenus Siphateles, 
although the simple intestine again is more like 
that of Rhinicluhys. The distinctive karyotype 
(see below) is most closely approached, among 
western minnows thus far karyotyped, by that of 
Gila (Siphateles) bicolor. 

From these observations and from geographical 
and paleohydrographic considerations, we con- 
sider it probable that Relictiis is an old relict 
( likely pre-Pliocene ) that was derived from an 
ancestral line from which the groups we refer to 
Gila and Rhinichthys were both also derived. 
Among living species of Gila, the generalized 
Utah chub {Gila atraria). despite its consistently 
biserial dentition, is probably a near relative, 
along with Gila ( SiphiUclcs ) bicolor. However, 
the combination of primitive and derived traits 
render the precise relationships of Relictiis diffi- 
cult to evaluate. 

Karyotype. Among the monotypic and poly- 
typic western genera of cyprinid fishes that 
logically may represent the closest relatives of 
Relictus (table 34), the chromosome number and 
morphology (figs. 47, 48 ) of the new genus stand 
distinctly apart (Uyeno and Miller, in prepara- 
tion). 

The karyotype is distinguished by a relatively 
large number of acrocentric chromosomes (2 
large and 8 small) but also many that are meta- 
centric (12). The remaining 28 chromosomes 
are subtclocentric and submetacentric, yielding a 
diploid number of 50, characteristic for all 
American cyprinids thus far examined (about 60 
species have been determined by Teruya Uyeno). 



Figure 46. Typical scales of adult females of three species of Cyprinidae from Nevada. A. Giki bicolor obesa 
(considered topotypic): 90 mm.; Humboldt River near Lovelock (G2); UMMZ 124873. B. Relictus solitarius (para- 
type): 90 mm.; Kirkpatrick Ranch. Butte Valley (Collection 7): UMMZ 141518. C. Rliiniclithys- <)«■!//;« robiistits: 
7.3 mm.; Indian Creek, tributary to Reese River; UMMZ 124894. Photographed by Louis P. Martonyi. 



194 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



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VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 195 




B 

Figure 47. Karyotype of Relictiis soliiariiis. processed hy Tcruya Uyeno in 1971. A. photograph of squash 
preparation. B, tracing of chromosomes, with some displacement where overlapping. The chromosome categories are 
analyzed and grouped in figure 48. 

Figure 48. Idiogram of chromosomes of RcIuiks si>liuiiiiis. Derived from figure 47. 






yi* 



196 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



In the idiogram (fig. 48). the 12 chromosomes 
interpreted as metacentric arc set apart in the 
first row, followed in the next two rows by the 
28 that are more or less transitional between 
metacentric and acrocentric, then, in the last row, 
by the 10. 2 large and 8 small, that are classed as 
acrocentric. Among the western genera men- 
tioned above, the karyotype of Rclictiis is per- 
haps most closely approached by that of Gila 
{Siplnilc'les) bicoh>i\ which also has 2 large but 
6 small acrocentric chromosomes and 16 meta- 
centric ones. Gila ( Gila ) robiisla has only 2 large 
and 4 small acrocentric chromosomes and 12 
metacentric ones. Rhiniciithys resembles Rclictiis 
in having 6 or 8 acrocentric but only 6 meta- 
centric chromosomes (Uyeno and Miller, in 
preparation ) . 

Derivation of name. The Latin rclictiis is 
the past participle of rcliqiio. to leave behind, and 
is also a noun, meaning a forsaking or abandon- 
ing. It is interpreted as a substantive, as in the 
English "relict." in the sense of the relict one. The 
gender is masculine. 

Relict Dace 

Relictus solitaiius Hubbs and Miller. 
(Figures 35C,D, 51.) 

Rcli<lus sdlitciiius HcHHS and Mil ilk. 1072. p. 102 
( diagnosis). 

Holotype. UMMZ 186904. a nuptial male 
60..^ mm. in standard length (fig. }5C). Para- 
types, all other .specimens (170. 34-99 mm. long) 
from same Collection (no. 7: data given below), 
including the mature female. 89.8 mm. long, that 
is illustrated (fig. 35D). All other specimens 
studied are listed below with an account of the 
status of the populations, but no others are 
designated as types. 

The discovery of this unique relict species is 
recounted under the heading of the genus. It 
occurs only in the contiguous drainage basins of 
the following pluvial lakes just south of the con- 
joining parts of the Lahontan and Bonneville 
basins, in eastein Nevada: Lake Franklin and 



its tributary Lake Gale, in Ruby and Butte val- 
leys, respectively, where native; Lake Waring and 
the tributary Lake Steptoe. in Goshute and Step- 
toe valleys, respectively, where also native; and 
Lake Spring, in Spring Valley, where almost 
surely introduced. The paleohydrography and 
the remnant waters of these ancient lakes are 
treated above (pp. 38-58). The evidence now 
seems conclusive that Rclictiis solilariiis is the 
only native fish in the expanse of 14,682 square 
kilometers that drained indirectly and directly 
into Lake Franklin and Lake Waring. It is prob- 
able that the species originally was relict in only 
one of the two main drainage complexes, but 
gained access to the other by stream wandering 
over a bajada across the low intervening Goshute 
Pass ( p. 43 ) . 

Within the entire north-central part of the 
Great Basin herein under treatment, this is by far 
the most numerous of the four native fish species, 
and it occupies about as many of the isolated 
spring waters as harbor the other three. It occurs 
in isolated springs and in springfed streamlets. 

The at least temporary establishment of this 
species in Spring Valley, to which we now as- 
sume it was not native, we attribute (p. 235) to 
its introduction from one of the north-central 
Great Basin valleys in which it is native. There is 
.some indication, furthermore, that it was once 
stocked in a spring in Utah ( p. 226). 

Material Examined and Population Status. 

Our Collections (table 35), with .some supple- 
mentation (p. 205). have provided a vast abun- 
dance of preserved material: 7.501 specimens in 
34 Collections, of which only 7 essentially dupli- 
cated a prior Collection. These Collections appear 
to represent the entire range of the species. In 
1971 a few additional specimens from previously 
sampled springs were taken, for identification and 
for their bearing on survival. 

We list the data in some detail, for the bearing 
they have on the ecology of this relict fish and 
on the population changes that have occurred, or 



VOL. VII HUBBS. MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



197 



Table 35. Data on collections of Relictus solitarius. 



Coll. 




Field 






No. 


Size, 


Map: 




No. Location 




No. 


Museum 


Cat. No. 


specimens 


mm. 


fig. 




Lake Franklin drainage 


















Ruby Valley 


















1 Borrow pit 




Lewis 4 


UMMZ 


186898 


70 


18-30 


8 




2 Pothole spring 




M 34-209 


UMMZ 


132188 


317 


14-59 


8 




3 Isolated spring 




Lewis 5 


UMMZ 


186899 


43 


17-50 


8 




4 Slough below Cave Cr. 


M 34-210 


UMMZ 


132189 


18 


24-39 


8 




5 Isolated spring 




Lewis 3 


UMMZ 


186901 


10 


10-17 


8 




6 Isolated spring 




Lewis 1 


UMMZ 


1 86900 


5 


24-27 


8 




Butte Valley 








(141518 


170 


^4-99) 






7 Kirkpatrick Ranch 


1 


H 42-47 


UMMZ 


(186904' 
132191 


1 


60 1 


8. 12 




8 Atwood Ranch ( = 


= 7) 


M 34-212 


UMMZ 


277 


21-99 


8, 12 




9 Atwood Ranch 




JED 64-53 


UNLV 




35 


21-84 


8. 12 




10 Trib. Butte Cr. 




M 34-211 


UMMZ 


132190 


225 


14-98 


8, 12 




1 1 Head. Odgers Cr. 




H 42-46 


UMMZ 


141517 


205 


24-77 


8. 12. 


14 


Lake Gale drainage 


















Butte Valley 


















12 Stratton Ranch 




JED 64-52 


UNLV 




67 


19-81 


8. 12, 


14 


13 Wrights Spring 




H 42-45 


UMMZ 


141516 


97 


27-64 


8. 12, 


14 


14 Stratton Ranch 




H 42-44 


UMMZ 


141515 


383 


11-72 


8. 12. 


14 


15 Owens Ranch 




H 42-43 


UMMZ 


141513 


219 


25-72 


8, 12. 


14 


Lake Waring drainage 


















Goshute Valley 


















16 Johnson Ranch (meadow) 


H 42-42 


UMMZ 


141511-12 


171 


7-99 


12 




17 Johnson Ranch (slope) 


H 42-41 


UMMZ 


141509-10 


706 


16-114 


12 




18 Phalan Creek 




M 38-165 


UMMZ 


124966 


168 


26-93 


8. 12 




19 S. ofCurrie 




M 38-164 


UMMZ 


124963 


201 


23-87 


8. 12. 


14 


Lake Steptoe drainage 


















20 Cardano Ranch 




M 38-162 


UMMZ 


124962 


257 


17-93 


8, 12, 


14 


2 1 Warm Springs Sta 


tion 


JED 62-27. pt. 


UNLV 


F 175 


147 


12-42 


8. 14 




22 Campbell Ranch 




M 38-166 


UMMZ 


124967-68 


341 


15-87 


8. 14 




23 Steptoe Ranch ( = 


22) 


JED 62-27. pt. 


UNLV 


F 174 


150 


13-41 


8. 14 




24 Grass Springs 




JED 62-27, pt. 


UNLV 


F 169 


155 


21-63 


8. 14 




25 Dairy Ranch 




M 38-160 


UMMZ 


124956-57 


367 


13-81 


8. 14 




26 Georgetown Ranci 


h 


M 38-158 


UMMZ 


124954 


403 


1 5-8 1 


8. 14 




27 Ruth Pond 




— 




— 


6 


— 


8, 14 




28 3C Ranch 




M 38-159 


UMMZ 


124955 


209 


17-89 


14 




29 Fish Pond Springs 


(=28) 


M 69-15 


UMMZ 


188959 


289 


20-82 


14 




Lake Spring (introduced) 


















30 Springs. Spring Va 


illey Cr. 


JFD 64-43 


UNLV 




80 


27-67 


12, 14 




31 Springs. Spring Vr 


dley Cr. 


JED 64-56 


UNLV 




99 


16-66 


12, 14 




32 Stone House 




M 38-48 


UMMZ 


124786 


1.073 


14-88 


12, 14 




33 Stone House 




M 59-90 


UMMZ 


177095 


498 


14-81 


12, 14 




34 Keegan Ranch 




M 59-89 


UMMZ 


177094 


39 


10-22 


14 





^ Holotype. 



may be anticipated, as a result of huinan activities. 
Locations of springs are specified so as to permit 
relocation for future checking. Evidence that we 
have gathered concerning the population status 
of the species is presented in this section. 

The Collections are serially listed under valleys 
(all in Nevada), from west to east, and within 
each valley from north to south. For this species 



we list Collections separately, even when they 
duplicated a previous sampling. 

Populations of Ruby Valley in former 

DRAINAGE BASIN OF PLUVIAL LAKE FrANKLIN. 

From Ruby Valley, which in pluvial times was 
largely occupied by the main body of Lake Frank- 
lin (pp. 39-45). we have examined 463 speci- 



198 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



mens from six Collections (1-6), which seem- 
ingly represent me-)st of the Recent range in the 
valley of this species — and of all native fish life 
in the basin, unless, as seems highly improbable, 
trout and/or chubs (Gihi hicoloi) were native 
in the mountain streams and in Ruby Lake, re- 
spectively. 

It is virtually certain that the relict dace was 
native in Ruby Valley, for King ( l<S78, p. 504, 
here quoted on p. 44 ) mentioned the occurrence 
of fish, presumably of this species, in Ruby Lake. 
On September I2-I.\ 1934, when he first ex- 
plored the basin. Hubbs found the dace abundant 
in the springs and sloughs of Ruby Lake, which 
occupied much of what had been the southern 
arm of pluvial Lake Franklin. Close to Collec- 
tion 2, from a single spring, the dace was found 
to be common on the gently sloping grassy bed of 
pluvial Lake Franklin in other waters examined: 
( I ) in other pothole springs, including one about 
2 m. in diameter and covered with 'blanket mos.s" 
(in these other springs numerous young about 
1 5-30 mm. long indicated late spawning) ; ( 2 ) in 
the rather swift, Nasturtiiim-contamxng outlet of 
a large stony seepage spring; and (3) in a large 
slough about 25 cm. deep within a large rushy 
marsh, which, with the sloughs along its western 
edge, constituted the bed of Ruby Lake ( this 
marsh contained many ducks, which would have 
been hunted later in the fall). Fish were lacking 
in icy-cold Cave Oeek (both within and below 
the cave) and in several large cold springs arising 
nearby from the same limestone formation. The 
fish was well known to residents as the "tule min- 
now" (a vernacular that was appropriate locally, 
but definitely is not applicable throughout most 
of the range of the species). 

A long-time rancher, W. J. Gardner, whose 
home ranch was on Gardner Creek, a tributary to 
the present Franklin Lake, reported to us in 1934 
that on the east side of Ruby Lake he knew of 
minnows occurring only, and intermittently, in a 
small spring three-fourths of a mile from Minnie 
Spring, which is southwest of the hot springs near 
the north end of the lake, but that Minnie Spring 



(presumably not named for minnows) was fish- 
less ( however, they were reported to be there 
later, by Donald E. Lewis). In 1932 Gardner 
had found "minnows" in a slough near his home 
ranch, after Franklin Lake, usually dry, had 
water; he thought these had come out of a tribu- 
tary canyon, but that they did not seem to be 
young trout. 

During our 1942 trip two ranchers in north- 
eastern Nevada testified regarding fish in Frank- 
lin River, the main stream course in the northern 
part of Ruby Valley leading toward Franklin 
Lake. One told of seeing small fish in the stream 
about 1937, but another thought it to be fishless. 
becau.se of its tendency to dry up ( we had found 
it dry in 1934). On June 28, 1942, Hubbs ex- 
amined the then flowing stream closely for about 
one-fourth mile on either side of the Ruby Lake 
Road, without locating a fish. 

Clearly, as late as 1934, the relict dace 
abounded in the profuse spring-fed waters of the 
Ruby Lake complex, but it is not certain that it 
iiccurrcd elsewhere in Ruby Valley. 

In 1965, Hubbs found that the dace had seem- 
ingly been extirpated in at least much of Ruby 
Valley, presumably as a result of the extensive 
development of a sport fishery, principally for 
largemouth bass (Micioplcnis salmoidcs), pro- 
moted by popular posters and leaflets and serviced 
by two .special landings with ramps for boats. 
The Nevada Fish and Game Department was 
operating a fish hatchery within the limits of the 
Ruby Lake National Wildlife Refuge (which also 
manages a public shooting area within the 
Refuge). The late J. Clark Salyer, head of the 
federal wildlife refuges, established the policy of 
protecting the endemic minnow by such means as 
prohibiting importation of bait minnows, but 
under the management practices of 1965 the 
minnow seemed doomed in Ruby Valley. In fact, 
not a single minnow was seen in that year during 
an examination of ten clear-water springs of 
various types in the long file emerging along the 
west edge of the basin, very close to the boundary 
between the Ruby Mountains bajada and the 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



199 



marshland representing the floor of Ruby Lake, 
from just south of Shanty Town for more than 4.8 
km. to one of the Ramirez Springs (all in T. 26 
N.,R. 57E.). 

However, at our request, Donald E. Lewis, 
Manager of the Ruby Lake National Wildlife 
Refuge, had his staff search for minnows through- 
out the refuge on August 2-14, 1967. This search 
led to the collection of Relictiis at four places, as 
is detailed in the following list. Mr. Lewis re- 
ported: 

The small fish are most abundant in isolated 
water where the largemoiith hass cannot reach 
them. However, I believe the fish does exist 
throughout the lake, but is greatly reduced by 
largemouth predation. All fish samples except 
No. 4 [Collection 1] were taken from isolated 
springs. No. 4 is an isolated dike borrowpit 
without a bass population. 

Mr. Lewis had earlier reported ( personal com- 
munication, November 12, 1965) having seen 
minnows about Ruby Lake at the North Sump 
(m NW '4 Sec. 5. T. 27 N., R. 58 E. ) on the 
west side of the lake; along the CCC Dike running 
across the lake near the northern edge of Sec. 22, 
in same township; at Minnie Spring, near center 
of Sec. 10, in same township (where earlier re- 
ported absent ) ; and at one place, where some 
were later sampled (Collection 5). He indicated 
that minnows do not occur in the Hot Springs 
(Sec. 2. T. 27 N., R. 58 E.), about 1 mile north- 
east of Minnie Spring. 

Collection 1 (fig. 8). Isolated borrow pit on border 
of flat of Ruby Lake, O.S km. east of western border 
of Wildlife Refuge, north of middle of Sec. 14, T. 27 
N., R. ."^S E.; Elko County, ca. 12 km. north of White 
Pine CoLmty. Donald E. Lewis, August 10, 1967 
(Sample no. 4); UMMZ I8689S (70, 18-30 mm.). 

Collection 2 (fig. 8). Pothole spring on gentle grassy 
slope on old lake bed, on western side of Ruby Lake. 
ca. 10 km. south of Harrison Pass, at elevation of ca. 
6,000 feet (1,829 m.): in or near Sec. 19, T. 27 N., R. 
58 E.: Elko County. Moderately clear water (bottom 
visibility ca. L.'i m.); soft mud, with some stones and 
slabs; slight outflow; much Chiini. Ncisturliiiin. and 
rushes — generally thick — but open in deep holes; cold. 
Hubbs family, September 12, 19.^4 (M34-209); UMMZ 



1321 88 (317, 14-58 mm.); 6-foot woven-mesh seine 
and 15-foot seine with '4 -inch square mesh. 

Collection 3 (fig. 8). Isolated spring between county 
road and the Ruby Lake flat 0.7 km. east of western 
boundary of Wildlife Refuge, .Sec. line 18-19, T. 27 N., 
R. 58 E.; Elko County, 10.5 km. north of White Pine 
County. Lewis. August 14, 1967 (Sample no. 5); 
UMMZ 186899 (43, 17-50 mm.). 

Collection 4 ( fig. 8 ) . Slough below Cave Creek on 
west side of Ruby Lake, in T. 27 N. near western border, 
about middle of R. 58 E.; Elko County. Moderately 
clear water; very soft bottom (collector sank 0.3-0.6 m. 
into the mixture of clay and sand); no apparent cur- 
rent; little vegetation (rushes and grass); moderate 
temperature. Hubbs family, September 13, 1934 (M34- 
210); UMMZ 132189 (18. 24-39 mm.); 6-foot woven- 
mesh seine. 

Collection 5 (fig. 8). Isolated spring near western 
border of Wildlife Refuge, just above lake flat, 0.3 km. 
south of northern border of T. 26 N., at R. 57-58 E. 
border; Elko County, 4.5 km. north of White Pine 
County. Lewis, August 9, 1967 (Sample no. 3); UMMZ 
186901 (10, 10-17 mm.). 

Collection 6 (fig. 8). Isolated spring 0.4 km. south 
of southern boundary of Wildlife Refuge, in NE 'A 
Sec. 12, T. 25 N., R. 57 E.; White Pine County, 7 km. 
south of Elko County. Lewis. August 2, 1967 (Sample 
no. 1 ); UMMZ 186900 (5. 24-27 mm.). 

In addition, Mr. Lewis provided one sample 
( R3 ) of a presumably introduced minnow, 
Rltinichthys osculiis rohiisttis (pp. 107-109). 

Populations of that part oh Butte Val- 
ley FORMERLY DIRECTLY TRIBUTARY TO PLUVIAL 

Lake Franklin. Five Collections (7-11), com- 
prising 9 1 3 specimens, adequately represent 
Ri'liciiis solitariiis in the northern part of Butte 
Valley, Nevada, that in pluvial time drained di- 
rectly into Butte Bay of Lake Franklin. Included 
is the type series of the species. Southward this 
area grades into the similarly spring-rich, fish- 
inhabited northern part of the section of Butte 
Valley that drained into pluvial Lake Gale. The 
whole valley to the north and west of Kirkpatrick 
( Atwood, Don Phalan) Ranch (Collections 7-9) 
appears from our field reconnaissance, from 
maps, and from local testimony, to be devoid of 
adequate habitat and of fish life. The informants 
included Jerome Phalan Stratton, White Pine 



200 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



6IE 



62E 



63E 



64 E 



30N 



29N 



28N 



27N 



26 N 




40 "30 



40°I5' 



25N 



40°00' 
II5°00' II4"'45' 

Figure 4^>. Sprinas. spring crocks and nicadovvs, intermittcnl streams and v\ ashes, and coLinty and township lines, 
in the spring-hcarnig. northern part of Bullc Valley, and adjacent regions, in FIko and White Pine counties, Nevada. 
Based on U.S. Geological Survey National Topographic Map, I :25(),0()() series, Elko Sheet ( I9?8), and Map of White 
Pine County, hy Fd. Millard & Son, Fly, Nevada ( l>-)3U); and on field reconnarssance by authors. Numerals refer to 
Collections of RclU ms s<>litaiiii\ (see pp. 1 ^'7-202 ) . 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



201 



County Road Supervisor and owner of Stratton 
Ranch. 

The hydrography of both drainages of Butte 
VaHey (fig. 49) has been described by Clancy 
(1968). 

Collection 7 (figs. 8, 12, 49). Ditch from upper, hill- 
side spring on Kirkpatrick Ranch (in 19.H called "At- 
wood Ranch," later called "Don Phalan Ranch"), lowest 
and northernmost spring water in valley, on east side 
north of the narrows, where opening onto lake flat in 
east part of T. 29 N., R. 62 E. (same location as for next 
two entries): Elko County, ca. 21 km. northwest of 
Currie. Clear water of fine taste; rather firm clay: rather 
slow current; little Polaiuogeton, cf. P. peclinalits; 18° 
C. (air 21°). Hubhs family. June 27. 1942 (H42-47): 
UMMZ 186904 (holotype, 60..^ mm.) and UMMZ 
141518 (170 paratypes, 34-99 mm.); one haul of a 6- 
foot woven-mesh seine. 

Collection 8 (figs. 8, 12, 49). Springs on flat area 
of Atwood Ranch (later "Kirkpatrick." then "Don 
Phalan" Ranch: same location as for Collection 7). 
Clear water: very soft false bottom at depth of ca. 2.0 
m.: moderate outflow; nearly choked with Cham. Utiic- 
iildiia, Myriophyllnin. etc.; moderate temperature. 
Hubbs family, September 1.^, 1934 (M34-212): UMMZ 
132191 (277. 21-99 mm.); 25-foot seine with U-inch 
square mesh in bag. 

Collection 9 (figs. 8. 12. 49). Spring on same ranch 
as for Collections 7 and 8. Clear water; gravel and mud; 
depth to ca. I m.; dense vegetation: abundant aquatic 
invertebrates; 18° C. (air 21°); J. E. Deacon and M. B. 
Rheuben, .September 14. 1964 (JED 64-53); UNLV 
(35, 21-84 mm.); 15-foot seine with ' 4 -inch square 
mesh. The spring source had been dredged out (banks 
as high as ca. 3 m.). 

Collection 10 (figs. 8. 12. 49). Western tributary of 
Butte Creek, fed by springs on property of Silver Butte 
Mining Co. (formerly "QuiHci Ranch," but known as 
"Dclker Spring" in 1942): north of narrows of valley, 
in northeast part of T. 28 N.. R. 61 E.; Elko County. 
Clear water: moderately soft mud; moderate current; 
choked with Naslurtiuiu. algae, etc.: moderate tempera- 
ture (warmer in summer). Hubbs family, September 13, 
1934 (M34-2I1); UMMZ 132190 (225, 14-98 mm.); 
6-foot woven-mesh seine. 

Collection 10 was made in the first spring 
north (ca. 2.4 km.) of Taylor Ranch (then a 
division of Odgers Ranch), on which the colder 
spring and its swift outlet, which was about 1 .5 m. 
wide and irrigated a large meadow, seemed to be 



fishless (residents thought that clearing, damming, 
and flushing of this spring creek may have killed 
minnows formerly present). This spring is prob- 
ably the one ("28/61-1 Idl") mapped and treated 
by Clancy ( 1 968 ) and shown on the cover illus- 
tration of his report. According to testimony of a 
former resident, springs at Odgers Home Ranch, 
ca. 8 km. .south of Collection 10 (now in Ruby 
Valley Indian Reservation), were also fishless. 

Collection II (figs. 8. 12, 14, 49). Head of Odgers 
Creek (headwater of Butte Creek). 0.8 to 1.2 km. below 
source (.Sheep Corral Springs, ca. 1.6 km. north of the 
then fishless Twin Springs), in Sec. 28 and 29. T. 27 N., 
R. 62 E.; Elko County, ca. 7.2 km. north of White Pine 
County. Very clear water, but very easily roiled to dense 
chalky-white; rather firm clay and gravel; swift, with 
pools; moderate vegetation (dwarf Cham. moss, etc.); 
12° C. at source (air 16°). Hubbs family. June 27. 
1942 (H42-46); UMMZ 141517 (205, 24-77 mm.); 6- 
foot woven-mesh seine. 

Populations of that i'art of Butte Val- 
ley FORMERLY TRIBUTARY TO PLUVIAL LaKE 

Gale. Four Collections (12-15), comprising 
766 specimens, well represent the species in the 
spring-water area, about 1 1 km. long, at the 
north end of the part of Butte Valley. Nevada, 
that drained into pluvial Lake Gale (thence into 
Lake Franklin). This area probably comprises 
the total habitat of native fi.sh within the Lake 
Gale drainage. This conclusion stems not only 
from our field work, but also from the direct and 
definite testimony of Jerome Phalan Stratton 
( mentioned above ) . who specified as fishless 
Snow Creek on Gibson Ranch ( in T. 25 S., R. 62 
E. ) and Youngs Ranch, both in the very arid 
southern .section of Butte Valley. 

Collection 12 (figs. 8. 12. 14.441. Spring on Stratton 
Ranch (see Collection 14); Elko County. 1.6 km. north 
of White Pine County. Sandy to muddv bottom; depth 
to ca. 1 m.; moderate current; dense vegetation; 18° C. 
(air 15.5°). J. E. Deacon and M. B. Rheuben. Sep- 
tember 13. 1964 (JED 64-52); UNLV (67. 19-81 
mm. ). 

Collection 13 (figs. 8, 12. 14. 49). Wrights .Spring, 
ca. 1.6 km. west of Stratton Ranch, in same township 
(T. 26 N.. R. 62 E.); barely on Elko County side of 
Elko-White Pine countv border. Moderatelv clear water 



20: 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



bill Ciisily roiled lo whitish; rather soil clay-mud: slight 
to moderate current; some Pouiiiioiicion. ct. P. pcctiua- 
tii.s: IS' C. (air l.V'). Hubbs family. June 2(\ 1942 
(H42-4?); UMMZ 141 5 Id (97, 27-64 mm.); 6-foot 
woven-mesh seine. 

The minnows apparently occurred along only 
ca. 30 111. of thi.s streainlet, which then formed a 
wet meadow; the upper 0.4 km. (in Elko 
County), which was shallow, swift, and partly 
filled with Cluiru and Nasturtium, was apparently 
fishless, as at Owens Ranch I Collection 15 ). The 
discharges from Collections 13, 14. and 15 were 
reported to remain unctmnected. but it seemed 
probable that they are separated only by more 
or less meadow-like areas which could be flooded 
by occasional torrential rains. 

Collection 14 (figs. S, 12, 14, 49). .Spring creek, O.X 
km. below springs (Collection 12), on Stratlon Ranch, 
near center of \ . 26 N., R. 62 E.; White Pine County, 
adjacent to Elko County. Clear water, of good quality; 
firm soil; moderate current; bur-reeds in and along 
water; LS ■ C. (air l.V). Hubbs family, lune 26, 1942 
(H42-44); UMMZ 14l,sl5 ( .VS.^, 11-72 mm.); nearly 
all taken in one 2()-loot haul with a 6-foot woven-mesh 
seine; including some young taken in meadow about 1.6 
km. farther downstream where tlitch is broader and 
largely choked with Cliara, etc.; resident said young ap- 
peared here every year, although entire meadow freezes 
in winter, aiKi that minnows also occur in sump against 
hills to southwest. Carp [C'ypiiniis ciiipio) and, re- 
portedly, trout [Salino or StilvcHnns) also occLirred. 

Collection 1 .■=; (figs. S, 12. 14. 49). .Spring creek on 
Owens Ranch, .s km. southwest of .Stratton Ranch (Col- 
lections 12 and 14), in SVV U ol \ 26 N., R. 62 E.; 
White Pine County, ca. 2.4 km. south ol Elko County. 
Moderately clear water, but easily roiled to chalky-mud 
color, of good quality; soft lo firm alkali clay; moderate 
to slow current; much Poniiimt^flcn. cf. /'. pccliiuitiis. 
sedges, etc.; 16 C. (air 19 ). Hubbs family, lune 24, 
1942 (H42-43); LJMMZ 141513 (219, 25-72 mm.); 
6-foot woven-mesh seine. In the half kilometer between 
collection and head springs, the feeders, swift .md with 
Nii\tuniiiiii. were avoided h\ the fish. 

Populations of Goshutk Valley, in for- 
mer DRAINACiE BASIN OF PLUVIAL LaKE WARlNCi. 

For a number of reasons we find it rather arbitrary 
and unsatisfactory to separate Goshute and Step- 
toe valleys. Nevada, for the groupings of Col- 



lections of Kclictii.\. One reason is that the springs 
in the northern end of Steptoe Valley, as custom- 
arily mapped, drain into Goshute Valley. There 
is no complete agreement on where the line divid- 
ing the two valleys cros.ses the common graben 
( as noted on p, 43 ), 

We treat this common graben as comprising 
the basins of two pluvial lakes, Waritig and Step- 
toe, while realizing that in Pleistocene time the 
two basins constituted a hydrographic unit, and 
may still be hydrographically connected in ex- 
treme flood. 

Four Collections (16-19). comprising 1.246 
specimens, represent the populations of Rclictiis 
that were found to occupy the disjunct spring-fed 
streamlets of the northern part of the Goshute- 
Steptoe valley complex below the basin of pluvial 
Lake Steptoe. We have found evidence of the 
occurrence here of this dace (and of any native 
fish) at only three locations: (1) the many 
springs on Johnson Ranch, where two Collections 
were made, near the north end of the valley (at 
the north end of the range of the species); (2) 
Phalan Creek. 72 arid kilometers to the south- 
southwest, the western tributary of Nelson Creek, 
which is the watercourse of the southwestern arm 
of Goshute Valley: and (3) the springs and ditches 
just south of Currie. in the stream course formerly 
followed by the river that connected Lake Step- 
toe with Lake Waring, 8-16 km. southeast of 
Phalan Creek. 

It seems probable that we have samples from 
all of the presently fish-inhabited areas in the 
pluvial drainage basin of Lake Waring below the 
outlet of pluvial Lake Steptoe near Currie. The 
one other site shown as plausible on maps, that 
of Flower Lake ( local u.sage ) or Flowery Lake 
(as mapped), which is near the center of T. 33 
N.. R. 66 E.. on the old lake bed. appeared from 
an examination on June 23. 1942. to be fishless, 
and it was so reported by experienced local folk. 
Although in the spring runoff of 1934 the playa 
of Goshute Valley had by reports been covered 
by water, very little remained of "Flowery Lake": 
one spring-fed pond, covering about 12 ■ IS m. 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



203 



and 0.3 m. deep in center, was choked with Chara 
and other vegetation; and. ca. 0.4 Icni. north a 
chain of small knoll springs watered a wet 
meadow (see also p. 54). There are no indica- 
tions that any sizable lake has existed in the area 
of "Flowery Lake." The Gibbes' map of 1 873 
(p. 8) merely notes at its location "Spring with 
grass." 

We regard it as certain that Relictiis was native 
in this drainage area. We secured no historical 
confirmation for Johnson Ranch (Collections 16 
and 17). but did for the two other Collections 
( 18 and 19). 

Collection 16 (fig. 12). Meadow springs on Johnson 
Ranch, east and north of ranch house, near north end 
of valley, on west margin, ca. 8 km. south-southwest of 
Oasis, in .Sec. 29, T. 36 N., R. 66 E.; Elko County. 
Moderately clear water (bottom visibility about 1.5 m.); 
firm clay around bulrushes, to extremely soft in deeper 
water: slight outflow from each spring; some choked 
with, or margined by, vegetation (mostly rushes or 
Nasturtium; one with much Utriculaihr. some with 
Portiiitogeton, cf. P. pcctinaliis; much algae in others), 
but others more open; uniformly 16° C. in all large 
springs with flowing outlets, but 21° in one spring with- 
out discharge (air 18° in early morning). Hubbs family. 
June 22-23. 1942 (H42-42); UMMZ 141511-12 (171, 
7-99 mm.); dip-net, 6-foot woven-mesh seine, and 25- 
foot seine with l<4-inch square mesh in bag. 

Waste water from the springs on Johnson 
Ranch flowed ( as Hardy Creek ) several kilo- 
meters .southward in a gully, but by 1942 did not 
provide suitable habitat for Relictiis. The 
meadowy spring area was unusually extensive: in 
the main group, near ranch house, there were 57 
fenced springs and others not fenced, and about 
1.6 km. north there were 15 or 20 other springs. 
The outlet of the one alluvial-slope spring (Col- 
lection 17) flowed along the south margin of the 
meadow — a point pertinent to the finding that the 
fish from the meadow springs have on the aver- 
age deeper bodies and wider heads than those 
from the upper spring, although tho.se from both 
habitats reach an unusually large size for the 
species (p. 214). Originally, before modification 
by ranching, the springs here probably produced 
a large area presumably occupied by the relict 



dace in large numbers. The Gibbes' map of 1 873 
(p. 8) shows a cluster of "Large Springs" with 
outlet leading .south about 18 km., with label 
alongside noting "Good Grass" ( indication of a 
meadow, where stream course is now entrenched). 

Collection 17 (fig. 12). Alluvial-slope spring, just 
.south of house on Johnson Ranch, and just above 
meadow springs (Collection 16). Very clear water of 
excellent taste; firm to soft clay and sand; moderately 
fast current; much Potaiuot^cton, cf. P. pcctiinitus, and 
algae, and lined with Nastiirtimn: 21° C. (air in after- 
noon 27°). Hubbs family. June 22-23, 1942 (H42-41 ); 
UMMZ 141509-10 (706, 16-114 mm.); 6-foot woven- 
mesh seine, and 15-foot seine with Vi-inch mesh. Fish 
abounded (one two-quart jar was filled by one haul of 
the 15-foot seine). Some were exceptionally large (fig. 
51). 

Collection 18 (figs. 8, 12). Outlet ditch of large 
spring (mapped as "Twin Springs"), on edge of Phalan 
Creek 3.2 km. above (west of) house of Phalan Ranch 
("Phalen Creek Ranch" on many maps),^ ca. 11 km. 
northwest of Currie. near southeast corner of T. 29 N.. 
R. 63 E., Elko County. Phalan Creek is the upper, 
western tributary of Nelson Creek, the main watercourse 
of the southwestern arm of Goshute Valley. Clear water, 
but easily roiled; mud and a little gravel, generally firm, 
with some stony marl; slight to swift, mostly moderate 
current; generally dense vegetation iNci.\turtiiiiii. Chara. 
and Polamogctiin, cl. /'. pcctiuatiis): 21° C. (air 29°). 
Hubbs family, August 25, 1938 (M38-165); UMMZ 
124966 ( 168, 26-93 mm.); 6-foot woven-mesh seine. 

Mrs. Zubiri, whose other testimony is men- 
tioned on p. 206, reported in 1938 that she 
found minnows common here as a girl, which 
would have been about 19(^2 or 1903. Some 
years prior to our conversation she saw minnows 
at the site of Collection 19. 

Collection 19 (figs. S. 12. 14). Large springs in nar- 
rows of valley, about 3 km. south of Currie. and upper 
1.6 km. of outlet ditch toward Currie, on either side of 
the T. 27-28 N. boundary, at middle of R. 64 E. (ca. 
40° 15' N. lat., 114° 45' W. long.); Elko County. Clear 
water; sand, clay, peat, and gravel; slight to swift cur- 
rent: much Chara. Nasturtium, and Potamogeton: 19° 
C. (air 31°). Hubbs family, August 24, 1938 (M38- 
1 64 ) : U M MZ 1 24963 ( 20 1 . 23-87 mm. ) ; 6-foot woven- 
mesh seine. 



' Jerome Phalan Slratton. Road Supervisor of White Pine County, 
informed us at tily, on June 25. 1942. that the proper speUing is 
Phalan ("grandmother Phalan brought inc into the world"). 



204 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



These springs are at the junction of intermit- 
tent McDermitt Creek ("Williams Cr."" on some 
maps) with the lava-entrenched canyon through 
which ancient Lake Steptoe discharged into Lake 
Waring. 

Already in 1Q38 evidence existed of the deple- 
tion of Rcliiins in this area. The storekeeper at 
Currie reported that minnows were formerly com- 
mon from the spring meadows to the road bridge 
in town ( though not above the meadows ) ; and 
that years previous Mr. Zubiri had caught min- 
nows here in a burlap sack for bait. On Septem- 
ber 13. 1934. during a cold night, Hubbs on ex- 
amining the stream from Currie to and including 
the headwater springs and ponds located no min- 
nows, though presumably at least a few were 
present. In 1938 he found the chubs only in the 
upper 1.6 km. of the stream, chiefly in quieter, 
deeper, weedier sections retaining a semblance of 
natural condition. Obvious factors in the deple- 
tion had been the straightening of the formerly 
tortuous stream course, accelerating the current; 
repeated cleaning out of the aquatic vegetation; 
diversion of the stream into irrigation ditches 
( we saw trout dying as a result ) ; and stocking of 
rainbow and briiok troLit iSdliuo gainlncrii and 
Sulvelinii.s jonlincilis). many of which were seined. 

Populations of Steptoe Valley, in for- 
mer DRAINAGE HASIN OF PLUVIAL LaKE STEP- 

TOE. Ten Collections ( 20-29 ). comprising 2.324 
specimens, rather aciequately represent the pop- 
ulations of Rclictii.s occurring in Steptoe Valley. 
Nevada, above the stream connection between 
ancient lakes Steptoe and Waring. The stations 
cover a north-south space of about 100 km., prob- 
ably enclosing nearly all of the habitats in this 
valley for the relict dace. The hydrography of 
the basin has been treated by Eakin ct <//. ( 1 967 ) . 
The most plausible additional habitat suggested 
by examining the relatively recent topographic 
maps is the cluster of springs shown in the Wil- 
low Creek basin near the western base of the 
Egan Flange, but an examination in 1969 of these 
waters, as well as of other places in the drainage 



basin of pluvial Upper Lake Steptoe, disclosed 
no trace of native fish (pp. 59-60). 

Evidence that Relictiis has long existed in Step- 
toe Valley despite the introduction of trout and 
other exotic fishes and despite other vicissitudes 
of existence, and that it is almost certainly native 
here, has been provided, for most of the Collec- 
tions (21-26). by long-time residents, including 
John Yelland, who had come into Steptoe Valley 
in 1881 and, when still observant and well pre- 
served, was interviewed by us in Ely on August 
22. 1938. 

Collection 20 digs. X, 12. 14). .Spring runs on Car- 
dano Kanch, on west sitle of valley between towns of 
Cherry Creek and Carnc. in .Sec. 5. T. 2.S N.. R. 64 E.; 
White Pine County, about 7.2 km. south of Elko County. 
Water clear, but easily roiled; firm to rather soft clay, 
mud. and a little gravel, with some stony marl: current 
none to slight; generalK choked with Niistiirlii(i)i. Chara, 
l'iil<iiii<ii:<liin. cf. P. p<'( liiniliis. Hippiiiiis. and filamen- 
tous algae; 14 C. (air }} ). Hubbs family, August 24, 
ly.^N (M3S-lh2); UMMZ 124962 (2.^7, 17-9.^ mm.); 
(i-loot woven-mesh seine. Local testimony in 19.^S in- 
dicated that bass (presumably Micmplci ii\ saliiioides) 
had occurred in the pond on this ranch. 

Collection 21 (figs. S, 14). Slough at Warm Springs 
(railroad station), near iNUmte Neva Hot Springs, about 
.^7 km. north of Ely. in Sec. 2.S, \. 21 N.. R. 63 £., 
White Pine Comity. E'irm mud and gravel bottom, he- 
coming mud near banks; no ciurent; 21 C. J. E. 
Heacoii. K. Larsen. and .L Tener, June 27, 1962 (JED 
62-27. m part); UNLV 175 (147. 12-42 mm.). John 
Nellant! testified in 193S that minnows had been here 
since the early ISSiS's. 

Collection 22 (figs. S, 14). Spring and outlet on 
Campbell Ranch (Steptoe P. O. ). on west slope of valley 
32 km. north ol l:ly and U) km. northwest of McGill, 
m Sec. .s. I. l'» N.. R. 6.^ E.. White Pine County. Clear 
water; nuid. gravel, ledges of spring deposits, etc.: slight 
to swift current: generally choked with Nasturliiini, 
CliiiKi. etc.: 24° C. (air 21). Hubbs lamily, August 
2S. I').^.S (IVI38-I66); UMMZ 124967-68 (341, 15-87 
mm); d-foot woven-niesh seine. Many carp (Cypiiinis 
iiiipi(i). and goldfish ( C((ra.s.si/r,s <iii)iilii\) . and some 
hvbrids between them, were also caught. The head 
reservoir, about 7 15 m., had just been washed out. 

Mr. Campbell, then 37 years old, the son of 
Ihc man who settled the ranch in 1878, felt sure 
that the minnows are native on the ranch, and 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



205 



other "old-timers" confirmed the occurrence of 
minnows here long ago. Campbell claimed that 
before the great smelter at McGill modified con- 
ditions deleteriously. the minnows abounded in 
the "Slough" (mapped as Duck Cr. ) below the 
ranch, to the extent that wagon loads could be 
gathered when the holes went dry in summer. 
John Yelland confirmed the evidence that min- 
nows had been common on this ranch since the 
early 1880's. 

Collection 23 (figs. 8, 14). .Springs on .Steptoe Ranch 
(same location as for Collection 22). J. E. Deacon and 
party, June 27, 1962 (JED 62-27, in part); UNLV 174 
(150, 13-41 mm.). The springs still contained carp 
(Cypriiuis carpio) and goldfish (Caiassius aiiratiis) as 
well as the minnows, indicating that these exotic cyp- 
rinids had not excluded the Rcliciux population over a 
period of more than a quarter century. 

Collection 24 (figs. 8, 14). Grass Springs, on Lusetti 
Ranch, 0.4 km. north of ranch house, on western side 
of valley about 27 km. north of Ely, draining into a 
slough in the flood drainage of Duck Creek; in Sec. 20, 
T. 19 N., R. 63 E.; White Pine County. J. E. Deacon 
and party, June 27, 1962 (JED 62-27, in part); UNLV 
169 (155, 21-63 mm.). 

Collection 24 was from one of three separate 
springs inhabited by this dace. A fourth, larger 
spring supported only Sacramento perch (Archo- 
plites interruptiis) , but. according to Ellen Vallee. 
the owner's daughter, it formerly also held min- 
nows. 

Collection 25 (figs. 8, 14). Upper spring ditch on 
Dairy Ranch, just below McGill (below reservoir used as 
swimming pool), in Sec. 20, T. 18 N., R. 64 E.; White 
Pine County. Moderately clear water (bottom visibility 
more than I.O m.); gravel and mud, mostly rather firm; 
uniformly moderate current; rather thick bottom growth 
of Potainogeton, cf. P. pectinatus, and considerable 
floating algae on sides; 25° C. (air 32°). Hubbs family. 
August 23. 1938 (M38-160) ; UMMZ 124956-57 (367, 
13-81 mm.); 15-foot seine with 'a -inch square mesh. 

Goldfish (Caiassius aiiialus). originally very 
brightly and variably colored according to local 
testimony, but since planting almost totally re- 
verted to wild type, were common here. John 
Yelland informed us that goldfish had been pres- 



ent for many years, along with minnows, in a deep 
hole on the Dairy Ranch. 

Collection 26 (figs. 8, 14). Several small springs 
and a little creek on Georgetown Ranch, in meadow 
just north of railroad yards of East Ely; tributary to 
Murray (sometimes corrupted to "Murry") Creek (open 
sewer of Ely, used for irrigation on ranch); in Sec. 2, 
T. 16 N., R. 63 E.; White Pine County. Clear water; 
spongy bottom; minute pools and riffles; generally 
choked with Nuslurtiiim and Potainogeton. cf. P. pec- 
tinatus; water cool. Hubbs family and Earl Manguni 
(local game warden), August 22. 1938 (M38-I58); 
UMMZ 124954 (403, 15-81 mm.); 6-foot woven-mesh 
seine. The largest spring, on south side of railroad 
tracks, was reported to have harbored many of the 
minnows before it was cleaned out and cemented in. 

Collection 27 (figs. 8, 14 I. Ruth Pond, just west of 
Ruth, in T. 16 N. (near middle of north border), R. 
62 E. Collected about 1964 by Dale V. Lockard, of 
Nevada Fish and Game Department, Wheeler District, 
Ely (6 specimens, not measured, received from him 
March 29, 1965, and returned). Probably the dace had 
been introduced here, well above the usual valley- 
bottom habitats. 

Supplementary Collections near Ely. After the 
text and maps of this report had been readied 
for the press, two additional collections of the 
relict dace have come to our attention. These 
were collected by Donald R. Cain of the Ely 
District Office of the U. S. Bureau of Land Man- 
agement and Frank N. Dodge of the Nevada De- 
partment of Fish and Game. The specimens were 
submitted to Miller for identification. The speci- 
mens have been sent to Japan, through Dr. Teruyo 
Uyeno. The first collection, of 10 specimens, of 
both sexes, 43-55 mm. long, was taken in Steptoe 
Creek 1 mile ( 1.6 km.) south of Ely, on May 24, 
1 97 1 . The second collection, of 10 specimens, of 
both se.xes, 36-7.3 mm. long, was collected at Fish 
Pond Springs, on C-B Ranch, at the same spot 
as Collections 28 and 29, described below. Mr. 
Cain rai.sed the question of the possible need for 
providing a sanctuary under Federal ownership 
for the protection of the fish at the C-B Ranch, 
in view of the danger to the fish imposed by 
irrigation practices and the removal of the aquatic 
vegetation. Miller had already discussed the 



206 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



question with James M. Vaughn at C-B Ranch 
in 1969. A sanctuary here for Relict us would 
indeed be a propitious prospect for the perpetua- 
tion of this unique endemic fish. 

On July 13, 1972. Robert J. Behnke and Frank 
N. Dodge collected 14 additional specimens, 31- 
75 mm. long: a few in Fish Pond Springs, where 
the dace were very scarce in the spring outflow, 
registering 13 C and the rest in a small stagnant 
pool, with murky water at ca. 27 C, and with 
tremendous numbers of dace showing no signs of 
stress. This pool, thought to be in the dry course 
of Steptoe Creek, was tiie same one seined by 
Cain and Dodge on May 24, 1971. 

We have recently learned that Thomas P. 
Lugaski, a student at University of Nevada, Reno, 
has collected specimens of Rcliclus soliUiriiis in 
Steptoe Valley and has been studying them. 

Collection 2S (tigs. S, 14). Springs on '}€" or 
"CCC" (Consolidated Copper Company) Ranch about 
1.6 km. north of ranch house, at base of truncated cones 
close to .Sleptoe Creek, atioul \i)5 km. south-southeast 
of Ely, near Sec. line 5-S, T. \5 N., R. 64 E.; White 
Pine County. Very clear water, of good taste; firm 
gravel to extremely soft organic mud; slight to moder- 
ately swift, occasionally swilt. current; generally choked 
with vegetation {Nasturliuni. Chain, and Poliiiin>i>clc>n, 
cf. P. pcdiniiliis): ') C. (air 27 ). Hiibhs family. Au- 
gust 2-\ l')3.S (M.^S-lsy); UMMZ \l-\'-)55 (209, 17-89 
mm.); (i-loot woven-niesh seine. 

Collection 29 (tigs. 8, 14). In same springs sampled 
by Collection 2.S; shown as Fish Pond Springs on Ely 
15-minule and Comins Lake 7.-S-niiniite quadrangles; 
ranch now named C-B Ranch. Very clear water, hut 
easily muddied; gravel, sand, and deep mud; slight ciu- 
rent; dense Chani. Ncisiiiiiiiini. and Potcimoiicu>n. cl. /-". 
pccthiatiis: ]'>' C. (air 27 ). Miller family, August 17. 
19h9 I M(i9-Ls ); UMMZ lSS9.su ( 2S(t, 20-79 mm., plus 
.^ skeletonized, 7()-S2 mm., plus some kept alive for 
chromosome study); 12-foot woven-mesh nylon seine. 
Clearly the relict dace had maintained ahLuuhmce here. 
A more recent Collection al the same place is men- 
tioned above. 

Additional evidence on the presence or 
absence of rlilictus at several localities 
IN Steptoe Valley not represented h\ col- 
lections. Cireen Ranch, west of middle of T. 



25 N., R. 64 E. Mrs. Zubiri (nee Green), a lady 
then about 50 years old, told us on August 23, 
I93K, that there never were minnows on this 
ranch except once, about 1902 or 1903, when 
hundreds, dead, appeared on the meadow below 
after a cloudburst. 

Murphy (formerly Dolan) Ranch, in south- 
western corner of T. 25 N., R. 64 E. Mrs. Zubiri 
added that as a girl she caught minnows in a sack 
in a pasture on this ranch, keeping "the larger 
ones, about 5 inches long, to eat like sardines." 
She said that the minnows then swarmed by the 
thousands in the springs. This statement almost 
surely reflects the former occurrence here of 
Relict IIS. although we collected only Gila atraria, 
obviously a recent introduction, probably with 
trout ( p. 58 ). It is further probable that manipu- 
lations to establish bass and trout, including the 
stocking of Utah chubs, have led to the depletion 
or elimination of Rclictus from this spring area, 
which provides a habitat seemingly normal for 
this endemic dace. 

Cherry Creek vicinity, in T. 23-24 N.. R. 62- 
63 E. This area does not appear to provide 
proper habitat for Rclictus. and local testimony 
in 1938 did not indicate presence of minnows. 
On June 26, 1942, Cherry Creek flowed inter- 
mittently near its mouth but contained no fish. 
Cherry Creek Hot Springs (45 C.). on alluvial 
slope 2.1 km. south of Cherry Creek (old town) 
fed a small reservoir cttntaining goldfish (Caras- 
siiis (iiinitus ) . 

Thirty-mile, Indian, and many other springs on 
route from Ely to Butte Valley. These are all 
mountain springs, small and surely without native 
fish, whereas practically all of the valley-bottom 
springs in Butte Valley contain minnows, "with 
no variation," according to Jerome Phalan Strat- 
ton and his ranch operator at Stratton Ranch 
(personal communications. June 25-26. 1942). 

Lackawanna Spring, on the western slope of 
valley 3.2 kiiL north of East Ely, in the northern 
part of T. 16 N., R. 63 E. This spring was said 
in 1938 by local residents William McGill and 
Earl Manguiu to have contained minnows [pre- 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



207 



sumably Relictiis] prior to the piping of the water 
into a bathhouse. 

Schell (often 'Shell') Creelv and vicinity, in the 
northern part of T. 22 N.. R. 64-65 E.. in the 
Schell (or 'Shell') Creek Range on the east side 
of this basin. Local testimony, and an examina- 
tion of Schell Creek on Scheilbourne Ranch 
(usually 'Shellbourne,' and variously otherwise 
spelled), a long-established and major enterprise, 
indicated in 1938 and 1942 that there were no 
fish in the creek or in mountain springs feeding 
this ranch, and that the only fish in the pond 
were goldfish, Carassiiis aiiratiis ( which John 
Yelland testified, in 1938, had been there for 
many years). This small pond, at the ranch, is 
fed by Lower Schell Creek Spring, which regis- 
tered a temperature of 26 C. in winter. Upper 
Schell Creek Spring, about '4 mile above, is cooler 
(22° C), and is diverted for domestic use. This 
information was furnished in 1971 and 1972 by 
W. E. Ireland and Robert L. Schultz of the 
Bureau of Land Management. 

The location during exploratory years of Schell 
Valley is discussed on pages 233-235, for its 
bearing on the original distribution of Rclictiis 
solitarius. 

Introduced Populations Of Spring Valley. 

In Former Drainage Basin Of 

Pluvial Lake Spring 

Five Collections (30-34). comprising 1,789 
specimens, represent the populations of Rclictus 
that we found in Spring Valley, Nevada, the site 
of pluvial Lake Spring. Some information perti- 
nent to the status of the populations is here in- 
cluded in the habitat descriptions, but the evi- 
dence that these populations have been derived 
through introduction is weighed in a subsequent 
section (pp. 233-235). The capture at Collec- 
tion 32 of three suckers, formerly thought to rep- 
resent an endemic species, but now definitely also 
attributed to an introduction, is treated below 
(pp. 229-230). 

Collection 30 (figs. 12, 14). .Springs adjacent to 
Spring Valley Creek (apparently a map name), near the 



northern end of the valley, at ranch about 10 km. north 
of Seigel Creek Road turnoff, in Sec. 31, T. 23 N., R. 
66 E.. White Pine County. Bottom of mud and organic 
ooze; cut hanks with sedges growing to edge. J. E. 
Deacon and party, June .5. 1964 (JED 64-43); UNLV 
(80, 27-67 mm.). Numerous deep springs here fed 
several canals that had been dug to a depth of about 
1.5 m. and contained abLmdant populations of the relict 
dace. About 0.8 km. north, similar springs were found 
to be fishless. Spring Valley Creek was said by rancher 
to contain fish when the flow was greater in the spring 
of the year (see Collections 32-33). 

Collection 31 (figs. 12. 14). Duplicate of Collection 
30, September \5, 1964 (JED 64-.*56); UNLV (99, 16- 
66 mm. ). 

Collection 32 (figs. 12, 14). Spring streamlet and 
pools above road crossing, arising in a meadow along 
the course of the mostly dry Spring Valley Creek, in 
the northern arm of valley, at Stone House (long a well 
known structure seen by the senior author in 1915 on 
the old Lincoln Highway between Scheilbourne and 
Tippetts, but by 1938 only seasonally occupied); in Sec. 
17. T. 22 N., R. 66 E.; White Pine County. Rather clear 
water (until disturbed); rather soft mud; current almost 
none to slight; dense Chura, surface algae, and other 
plants; 23° C. (air 24°). Hubbs family and Miller, July 
6, 1938 (M38-48); UMMZ 124786 (1,073, 14-88 
mm.); 6-foot woven-mesh seine and derris root in pool 
at bridge. 

Collection 33 (figs. 12. 14). Same spring. Clear 
water, but easily muddied; soft mud above firm soil; 
slight but definite current; dense vegetation (dense beds 
of Potamogelon. cl . P. pcclinatus. and green algae, with 
some Leinna and Hippiiiii.s. and a Sciipiis border); 22° 
C. (air 27 ). Hubbs and Miller families, July 5. 19.^9 
(M59-90); UMMZ 177095 (242. 14-81 mm.; plus 7 
skeletons; plus 249, 31-81 mm., in exchange series); 
15-foot seine with '4 -inch square mesh. 

Collection 34 (fig. 14). Ditch in spring-fed meadow 
about 0.8 km. east-southeast of the then abandoned 
Keegan Ranch house on the west side of valley flat in 
.Sec. 12, T. 18 N., R. 66 E.; White Pine County. Clear 
water; mud bottom; slight current; much vegetation; 
about 20° C. Miller and Hubbs families, July 5. 1959 
(M59-89); UMMZ 177094 (39. 10-22 mm.); dip-nets. 

The occurrence here of young dace in the 
meadow strip on the west side of Spring Valley 
appears to be attributable to wash-down from 
Spring Valley Creek, probably in one of the oc- 
casional floods ( on July 6, 1938, Bert Robison in- 
formed us of a flood in this stream course, follow- 
ing the drought of 1934. that had periled barns 



K18 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



E 100 
E 



SPECIMENS 

■ 706-1,073 
□ 201-198 

■ 117-171 
n 80-99 

■ 35-70 
D 5-18 



I 



-I 80 
Q 

cr 

< 

Q 70 

< 

<J^ 60 



Q_ 

oo 40r 



< 

PLUVIAL ! — 

LAKES 

VALLEYS 



n 



D 



D 



R A N K L I N 



n □ 



GALE 



2 3 4 5 eT^ 



B U^ T T E 

9 10 I ll 12 13 14 15 



D 



WARING 



D 



n 



n 



S T E P T E 



GOSHUTE-STEPTOE 



16 17 18 19 20 21 22 23 24 25 26 28 29 



D 



SPRING 



SPRING 



30 3 1 32 33 34 



COLLECTION NUMBERS FOR R E L I CTU 5 S0LITARIU5 
Figure 5U. Miixinium size of specimens in all Collections of Relicuis solirariiis. Data from table 35. 



nearby ) . A half-hour search for other popula- 
tions in springs and ditches for 0.4 km. below 
the ranch house disclosed no further populations, 
and an open, clear, spring-fed ditch below the 
ranch contained no fish. However, Phil Rowe. 
who owned the Bassett Ranch about 2.4 km. 
north, told us that he had .seen many small fish 
in the slough leading to the reservoir below where 
we collected, and that the fish appear in time of 
high water. We found the reservoir dry except 
for one muddy lake. 

Di:SCRlPTION AND COMPARISONS. 

Many of the more distinctive features of 
Ri'lictiis soliiaiiii.s are presented in the preceding 
account of the genus, along with a discussion of 
the relict status and relationships of the species. 

The characters of the species closely simulate 
those of other .spring-inhabiting fishes of the West, 
such as we have outlined on page 181. These 
spring isolates are cast into such a similar mold 



as to obscure their origin, relationships, and 
classification. As Hubbs and Miller ( 1948b, pp. 
51-52) indicated, this circumstance rendered 
dubious the generic placement of the species. This 
doubt persisted until we considered it generically 
distinct on osteological grounds. Even before that 
study was made, however, we thought that it prob- 
ably deserved generic separation from Rhinich- 
tliys. 

Sizi;. Relictii.s soUtarhis is a medium-sized 
minnow, ordinarily reaching a maximum standard 
length of 60-100 mm. (table 35: fig. 50), usu- 
ally larger than Rliinichlliy.s osciiliis but smaller 
than Gila hicolor. Only one of the 34 Collections, 
no. 17, one of the two at Johnson Ranch in 
Goshule Valley, contained fi.sh longer than 99 
mm., and only 5, among the total of 706 speci- 
mens taken there, were larger (101 to 114 mm. ), 
and they look abnormal (fig. 51) and are re- 
garded as superannuated (p. 214). Four other 
Collections barely missed reaching 100 mm. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



209 



(table 35; fig. 50). Specimens longer than 60 
mm. are included in all Collections, with the 
following exceptions: all Collections in Ruby 
Valley, where the species appears to have been 
dwarfed (our largest specimen measures 59 mm.); 
Collections 21, 23, and 34, for which circum- 
stances suggest that the larger adults were not 
taken; and Collection 27, which contains few 
specimens, not measured. At least one fish as 
long as 63 to 99 mm. is contained in 23 other 
Collections, and at least one longer than 70 mm. 
occurs in each of these 23 series that contain 
more than 200 fish, and such a size was attained 
in each valley, other than Ruby Valley, and in 
each pluvial drainage basin represented. There 
is little basis for thinking that markedly different 
sizes arc attained in any of the basins inhabited, 
other than Ruby Valley, and even there the re- 
duced size indicated for this species may be 
attributable to environmental conditions, rather 
than to an inborn characteristic. Predation may 
have been heavy at times, even primordially, be- 
cause the spring waters are relatively open and 
aquatic birds are common during migration. At 
present, predation by bass is so heavy that limited 
populations of the dace have survived only in 
isolated springs. Originally a larger size may have 
been attained than is indicated by our one main 
series (Collection 2), for, although local residents 
had observed only such small fish in the adjacent 
slough, larger ones, thought ( probably wrongly ) 
to have been of a different species, had been seen 
in certain springs. However, small specimens in 
Collection 2 had already developed the heavy 
pigmentation of the adult type, suggesting that 
indeed the Ruby Valley population was dwarfed, 
at least when sampled. There is a possibility, 
albeit seemingly very remote, that another larger 
minnow, most plausibly Gila bicolor. occurred 
in Ruby Lake prior to its elimination by bass. 

Coloration. The general color tone of half- 
grown and adult specimens in alcohol, save for 
the isolated blackened regenerated scales ( de- 
scribed below), is ordinarily rendered almost 
evenly dusky by innumerable melanophores that 



are arranged without a trace of the concentration 
around the scale-pocket margins that is usually 
evident in cyprinids ( this lack of scale margining 
may be attributed to the deep embedding of the 
scales). The melanophores become sparser ven- 
trally and in some adults ( including the holotype) 
remain undeveloped on the ventral surface of the 
trunk, whereas many other specimens are punc- 
ticulate there. 

The melanophores are of two sizes. Small ones 
are the more numerous, but large ones, as in the 
subgenus Sipluifeles, typically occur ventrally, and 
frequently predominate there; and some often oc- 
cur dorsally. The melanophores that occur ex- 
clusively or predominantly on the darkened re- 
generated scales (see below) are of the larger 
size. On the silvery opercle, the main part of the 
cheek, and the exposed part of the shoulder girdle, 
the larger chromatophores are conspicuous and 
the smaller ones are few. In contrast, minute 
melanophores render dusky the region bordering 
the eye, the entire top of the head including the 
snout, and also the suborbital region, where, oc- 
casionally, a concentration of pigment and a 
downward extension of the pigmented area simu- 
late a short, oblique subocular bar. The lower, 
more silvery part of the iris typically bears few, 
scattered large melanophores and few small ones, 
whereas the upper part is densely puncticulate 
with small ones. The dark upper and lower lips 
and the anterior intergular region become densely 
punctate in adults, especially in males, leaving 
largely clear most of the dentary, branchiostegals, 
posterior intergular, and exposed shoulder-girdle 
regions. The narial flaps are also puncticulate, 
especially toward the margin. The biting edges 
of the lips are clear of pigment, but the floor of 
the buccal cavity is darkened by melanophores 
which, in subadults, are few and are restricted to 
the forward part of the cavity. 

There is almost no trace of the lengthwise 
striping that is typically more or less sharply evi- 
dent in Rliinichthys. In the superficial pigmenta- 
tion there is no indication of a longitudinal streak 
on the head, such as Rhiiiicluhys characteristically 



210 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



displays, at least on the snout. Some specimens of 
Rclictiis, however, show a deep-lying dusky area, 
without obvious melanophores, below the nostril. 
Moreover, there is no longitudinal dark band on 
the body, except, in some specimens, for a deep- 
lying axial dusky streak, similarly without super- 
ficially discernible melanophores. This streak 
tends to be restricted to the posterior part of the 
body, and in some Collections, without sharp 
geographic significance, is rather conspicuous. A 
special concentration of the black regenerated 
scales along the midlateral region, presumably 
resulting from an injury, occasionally presents 
the impression of an irregular lateral band, or 
even, rarely, a row of short cross bars. There 
is tremendous variation in degree and type of 
speckling due to the irregular kiss and regenera- 
tion of scales. 

The fin bases and the middorsal and midventral 
lines in the adults are remarkably free from the 
pigmentary striping that is seen in most American 
cyprinids. Ordinarily, in Rclivliis. there is the 
barest, diffuse trace of a darker stripe from dorsal 
fin to occiput; no especially darkened or clear 
areas at the origin or midbase of dorsal fin ( some 
smaller adults and subadults do show a somewhat 
depigmented area about the front of dorsal and 
some darkening of base medially ) ; and no dorsal- 
midline intensification of the deep-dusky pigmen- 
tation between dorsal and caudal fins. Melano- 
phores are vaguely clustered posteriorly on the 
lower surface of the caudal peduncle, but do not 
form there a definitive band or streak, either 
superficial or deep. About and immediately be- 
fore the lower prociurent caudal rays there is a 
small clear area, which is noi matched at the ckir- 
sal edge. This dace therefore lacks the paired 
light areas at the caudal base that are ordinarily 
seen in Rhinicliihys. 

The fins in the adult ordinarily become dark- 
ened by numerous melanophores. which lend to 
be concentrated along the middle of the rays 
basally but along the margins of the branches of 
the rays distally. The chromatophores are fewer, 
and are retarded in development, on the pelvic 



and anal fins, where they are very few in many 
subadults and in half-grown fish, and may even 
be weak or in some localities absent, in some 
larger adults. These fins are largely clear mar- 
ginally. The pectoral fin is dark to the margin, 
except on the inner-ventral part. In develop- 
ment, the membranes of the dorsal fin, the caudal 
fin (especially on the lobes), and the pectoral fin 
near its front-upper edge, rapidly become 
crowded with small melanophores. whereas else- 
where the fin membranes remain largely clear. 
In these several respects, the mature males tend 
to assume the adult coloration more rapidly and 
more fully than do the females. 

The peculiar pattern referred to above, caused 
by irregularly scattered scales that have been 
blackened by being beset over the entire exposed 
smface by large melanophores. imparts a Rhin- 
/c7;//;y.s-like appearance to Rclictii.\, but is not a 
trustworthy indication of phyletic relationship. 
The same pattern (Langlois, 1929, p. 161 ; Hubbs, 
1 942, p. 5 ) occurs in other cyprinids. It is a con- 
spicuous feature of the several subspecies of the 
probably not closely related eastern North Ameri- 
can cyprinid now known as Seinotilus margaiita 
(Cope), which for some time previously was re- 
ferred to a monotypic genus Margarisciis. This 
pattern, occasionally rather conspicuous, but 
commonly barely evident, is also seen in another 
probably not intimately related cyprinid, Couesiiis 
plunibciis (Agassi/), which for some time re- 
cently was referred to Hyhopsis ( Bailey et al., 
1960. p. 14). but has now been referred again 
to Coiii'sius (by Bailey et al.. 1970. p. 19). The 
same pattern we find to hold for another probably 
distantly related cyprinid. the peculiar Pacific- 
drainage minnow Oregoiiiihthys cntincii (Sny- 
der), which was originally, and again recently 
( Bailey ct al. 1960. p. 14; Bailey et al.. 1970. p. 
20) referred to Hyhop.sis. but which, according 
to osteological research by Ted M. Cavender 
(personal communication. 1970). deserves sep- 
aration in the distinct genus Oregonichthy.s 
Schultz and Hubbs (1961); Reno (1969) has 
also suggested that Oregoiiichthys deserves generic 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



211 



status, on the basis of the unique structure of its 
neuromasts. (Similar appearing blackened scales 
that are conspicuous in two eastern species cur- 
rently referred to Hybopsis. namely H. (Erimy- 
st(i.\) dissimilis (Kirtland) and H. {E.) x- 
punctatiis Hubbs and Crowe (1956), are basic 
elements of the color pattern, unrelated to scale 
regeneration. ) 

The peritoneum is silvery, with brown punc- 
ticulations. 

The pigmentation develops very early. At a 
standard length of 10-18 mm., as indicated by 
Collections 1 and 5, the melanophores are already 
well developed. The middorsal streak has not yet 
developed, except along middle of dorsal base. 
where it is black. There is a similar streak along 
base of anal fin. A fine black streak has de- 
veloped along the a.xial septum of the lateral 
muscles. The dark blotch below the nostril has 
already become rather conspicuous, but does not 
form a lengthwise band, such as is seen in cor- 
responding young of Rhinichlhys. 

When a length of 20-25 mm. has been at- 
tained (as indicated by Collection 3), a dark, 
deep-lying, rather diffuse middorsal streak has 
appeared, more strongly before than behind the 
dorsal fin. It shows along the middle of the 
dorsal base, which, however, is pale about the 
front of the fin. No definite streak has formed 
behind the anal fin, where some separated 
melanophores, not evident in smaller fish, have 
formed. A file of 1-6, usually 2-4, huge melano- 
phores lie along each side of the anal base 
medially (these are inconspicuous when the pig- 
ment granules are centrally concentrated, and in 
some specimens they appear to be lacking). A 
deep-lying, diffuse axial streak has appeared about 
the now weak black axial line. A very irregular 
dorsolateral file of large melanophores has 
formed but soon fades. Little pigment has yet 
developed below the eye. 

Life color. The life colors of Rclictus 
soUtaiins were found to be extremely variable, 
perhaps in correlation with its secretive habits 
and with reduced selection pressure in its isolated 



habitats. The colors were described by us in the 
field for Collections 2. 7, 8, 10, 11, 15-18, 20, 
22, 25,26, 28, 32. and 33 (pp. 199-207 ). which 
represent well the geographical range of the 
species. The life-color variability is another 
unique feature of the relict dace. 

The colors are highly variable individually and 
to some extent geographically, but with con- 
siderable coherence in some respects. The gen- 
eral tone is usually dusky, strongly speckled with 
brown or, in some localities, with moss-green. In 
addition to rather bright blue reflections, which 
are generally apparent, there is more or less gilt, 
varying individually and regionally from almost 
none to quite bright. Many possess also a pale 
to deep tinge of violet (especially noted for Col- 
lection 2). Coupled with this, usually, is an ex- 
ceptionally heavy coating of slime. 

At some localities the general color was noted 
as especially variable. Thus, in Collection 32, the 
general tone graded from silvery gray, through 
pale to deep olive, to rather rosy (but never red) 
or to purplish olive-brown, with reflections from 
individual scales varying from silvery to bright 
blue or gilt. Some specimens (as noted for Col- 
lection 28 ) tended to grade toward yellow or 
gray. In some examples (as recorded for Col- 
lection 25), the entire back is rather uniformly 
pea-green and the mid-sides are either silvery or 
gilt; other specimens have the sides dappled with 
bright metallic-green .scales: some vary toward 
olive or gray. In others ( as observed in Collection 
17), the general tone is usually metallic greenish, 
bluish, or bras.sy; some fish (as in Collection 16) 
are even more variable — silvery, greenish, golden 
or, often, yellow-tan, and, on the average, defi- 
nitely more gilt than in some other series. In 
Collection 1 5 the various hues on the upper parts 
ranged from bright brassy green to dark olive or 
grayish green. The back (as it was in Collection 
10) may be bright grassy green. 

Generally the lower parts are lighter, often 
silvery or watery white; or silvery white, usually 
with a blue tinge; but in .some, either the dusky 
dorsal color ( in darker fish, as observed in Col- 



21 2 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



lection 18). or the yellowish color, extends over 
the ventral region. Intervening between the upper 
and lower sides is a narrow deep-lying dorsolateral 
stripe that is often pearly or golden ( as .seen in 
Collection 17) or bright brassy (Collection 10). 
The niidsides are often largely silvery, with more 
or less conspicuous reflections of bra.ssy or violet. 
In very large females (as in Collection 17) con- 
siderable gilt may show along a midventral streak 
on the abdomen, as well as on the side of the body 
and on the lower fins and their axils. 

The lower fins are often yellowish or lemon, 
occasionally rather bright golden or yellowish (as 
seen in Collection 32); watery white in others, 
especially in small females; but never red. The 
dorsal and caudal fins are usually more or less 
amber, often suffused with greenish or yellowish, 
either light or dusky; sometimes dusky gold. The 
lower fins show, at least in some fish, blackish 
speckling on the rays and broad blue-gray bor- 
ders. In some Collections ( as observed in Collec- 
tion 16), all fins may be yellowish, except near 
the margin. 

Many individuals are waxy or watery yellow- 
ish or lemon-colored on the pectoral and pelvic 
axils, but, unlike Rliinichfhys and some other 
western minnows, this species never has either 
axil red, even in breeding males. As observed in 
some C oUections. breeding males may be brighter 
than females, but none have red fins. Breeding 
males may have sooty lower fins (as seen in Col- 
lection .33). 

The opercle and the cheek behind the eye are 
more or less gilt, but there is no definite bright 
color bar at front of opercle. Very large speci- 
mens (as observed in Collection 18) may be 
bright blue across the upper edge of the opercle. 
The iris (as noted for Collection 28) may be 
golden only on a narrow inner rim. or (as seen in 
Collection 25 ) may be bright silver-gilt about 
pupil and sometimes gilt or orange elsewhere. 

One adult ( in Collection 25 ) was so golden 
that it appeared to be a mutant. Its back was a 
mixture of green and blackish; the sides were 
bright, clear golden yellow; the lower fins were 
a bright and clear, rather intense, golden yellow; 



the caudal fin was of the same color, with a large 
blackish blotch on lower lobe and finer blackish 
marking on the upper lobe. Some other speci- 
mens were sufficiently extreme to approach a 
mutant appearance. In addition to the blackish 
regenerated scales ( which are often concentrated 
near midsides or toward base of caudal fin to 
form seemingly aberrant color markings), jet- 
black blotches occasionally appeared on the body 
or fins (for example, on front edge of dorsal fin 
near base in an adult in the type series, Collec- 
tion 7). 

Form and general appearance. This is a 
rather chubby fish (fig. 35C,D), which the field 
notes indicate as relatively soft-bodied. However, 
it almost invariably preserves well, with very little 
tendency toward further softening or distortion. 
The deep imbedding of the scales imparts a fleshy 
feel to the surface, which tends to be slimy. 

The general form is that of a comparatively 
sluggish midwatei' swinnner. with rather deep 
body and deep peduncle, and rounded dorsal and 
ventral contours which are relatively symmetri- 
cal, with the dorsal profile not very much more 
curved than the ventral. The muzzle is rounded 
in both the lateral and the dorsal aspect and the 
rather large, straight, and oblique mouth rises 
forward, so that the upper lip, which is about 
even with the lower or only slightly protrudes, is 
about on the longitudinal axis of the body. It is 
not a conspicuously streamlined fish. The fins 
are relatively small and are generally strongly 
rounded; the paddle-shaped pelvics are particu- 
larly distinctive. The fins tend to be very flexible, 
often rather silky toward the margin. With age, 
the rays commonly branch more extensively than 
is usual, so that in many specimens they fan out 
at the margin to touch one another. 

Primarily because of the posterior location of 
the pelvic, the dorsal origin varies from slightly 
before to slightly behind the vertical from the 
pelvic insertion, usually slightly behind. In this 
respect Relictiis agrees essentially with Giln hi- 
color. but contrasts with Rhinicluhys. which has 
the origin of the dorsal fin behind that of the anal 
fin. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 213 




Figure 51. Relictiis solirwiiis: superannuated females from Johnson Ranch, Goshute Valley. Nevada; Collec- 
tion 17. A. UMMZ 141510, no. 2, 107 mm. B. UMMZ 141510, no. 4, 114 mm. C. UMMZ 141509, 113 mm. 



214 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



The general appearance of this fish contrasts 
particularly with that of Rhinichiliys. though it 
and sonic of the most modified of the spring- 
restricted forms of that genus converge consider- 
ably. 

Most of the features just mentioned seem to 
align this fish with Glhi hicolor, hut its duskier 
color, generally smaller and more imbedded 
scales, very incomplete lateral line, less regular 
squamation, much slimier integument, softer 
flesh, and generally smaller size belittle the re- 
semblance. 

In many of the aspects of form this .species, 
in comparison with other western minnows, 
seems to be unusually variable. Differences in 
form in addition to those that appear in compar- 
ing the measurements arc rather notable in several 
respects. In some specimens the upper profile of 
the head is more nearly horizontal and is much 
less decurved anteriorly than in others. This char- 
acter is usually associated with a large mouth and 
an especially turgid upper lip. but the correlation 
is by no means imiform, and this character is 
highly variable and is far from consistent at any 
one locality. This modification of the head pro- 
file is most strikingly evident in Collection 1 7. one 
of the two at Johnson Ranch, far north in Cioshute 
Valley, and is particularly pronounced in the 
oversized females, of which three are illustrated 
(fig. 51 ). Such modification is displayed to some 
extent in a varying proportion of the specimens 
in other Collections. There is also some variation 
(not measured) in the obliquity of the mouth. 
The width of the opercular membrane also 
fluctLiates considerably. 

The modification of form in the few definitely 
iwersized specimens in Collection 17 is extreme 
but not consistent. They differ among themselves 
in the robust build, great depth of peduncle, de- 
gree of nuchal hump, prominence of muzzle, etc. 
Such extreme modification of form is connnonly 
seen among oversized fish. Thus, an immense 
adult female of Gila orciitlii ( Eigenmann and 
Eigenmann) that came from a stream into which 
the species had recently been introduced in north- 
ern Santa Barbara County. California had a 



physiognomy much like that of these huge speci- 
mens of Relictiis. 

Lateral-line system. In this species, as in 
some other minnows inhabiting i.solated springs, 
the lateral line is greatly reduced, often inter- 
rupted, and irregular. It rarely extends to below 
the origin of the dorsal fin, and the part that is 
developed is often incomplete. In 90 subadult 
and adult specimens from various parts of the 
range of the species the pores per side range from 
.^ to 29, with 68 (76 percent) from II to 22, 
and with the mean at 16.02. The holotype has 
17-19 pores, ending 5-10 scale rows before verti- 
cal from origin of dorsal fin. 

On the head as well, the lateral-line system is 
deficient in some respects. The supratemporal 
canal is complete across the occiput in only 4 
among 76 specimens examined from various Col- 
lections: these four have 6-8 pores. In the 72 
specimens I 144 sides) found to have the canal 
medially interrupted (as often in spring-inhabiting 
isolates), including 27 specimens from Collec- 
tion 22 (not listed in table 4,^) that contain the 
fcHU" with the canal complete, the number of pores 
per lateral segment on either side ranges from 
to 5, with 92 counts of 3 and 34 counts of 4, 
and with a mean of 3.42 pores. The prcoperculo- 
mandibular pores in 222 counts (2 counts per 
fish ) range from 1 1 to 1 9, in a rather similar pat- 
tern in the several valleys; average. 13.79. The 
mandibular pores (table 44) in 158 counts (79 
specimens) range from 3 to 8, modally 5, with 
means fluctuating with the valley from 4.93 to 
5.61. 

Scale structuri;. The scales (fig. 46B) are 
more like those of Rhinichrhys than those of Gila. 
but usually bear even more numerous radii on all 
fields. Typically they are vertically oval, with 
the focus definitely nearer the base than the po.s- 
terior margin; but, with increasing size, the scales 
in some individuals ( sampled above lateral line 
just before dorsal fin) become longitudinally 
rectangular with dorsal and ventral edges .straight 
and horizontal, and with the focus even closer to 
the base. The two types integrade at a single 
locality. 



VOL. Vll HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



215 



Table 36. Proportional nicasiireiucnls. in pcruultat;e of standard leni^tli. f(ir Rcliclus solitarius in different pluvial- 
lake drainages. For each entry there is given the range, and below this the mean: niiiuhers of specimens for each 
category and the Collection numbers involved are shown in table 37. or. when fewer were counted, as a subscript. 



Predorsal length 



Anal origin to caudal base 



Sex 
S.L., mm. 



Males 



Females 



Males 



Females 



24-42(-) 42-66 30-42(-) 42-66(-) 66-114 24^2(-) 42-66 30-42(-) 42-66(-) 66-114 



Lake Franklin 
Ruby Valley 

Butte Valley 

Lake Waring 

Lake Steptoe 

Lake Spring 

Total range 
Grand average 

Lake Franklin 
Ruby Valley 

Butte Valley 

Lake Waring 

Lake Steptoe 

Lake Spring 

Total range 
Grand average 

Lake Franklin 
Ruby Valley 

Butte Valley 

Lake Waring 

Lake Steptoe 

Lake Spring 

Total range 
Grand average 



534-616 
581 

554-594 

573 

570-631 
600 

551-613 

571 

564-620 
593 

534-631 

585 



557-588 
574 

549^586 

577 

558-634 
594 

556-592 
571 

548-596 
578 

548-634 

585 



55()_6()7 566-599 



577 



587 



307-354 
329 



545-615 576-619 593-626 278-316 



580 



596 



604 



295 



589-628 581-642 594-688 286-353 



604 



611 



622 



326 



553-600 570-636 532-632 301-332 



581 



602 



589 



587-607 582-617 586-626 



595 



597 



607 



313 

309-339 
322 



545-628 566-642 532-688 278-354 



588 



602 



608 



Body depth 



259-290 241-276 265-301 258-291 
274 261 285 272 



320 



125-146 
137 



271-312 258-302 276-347 272-297 243-302 133-159 

290 281 290 283 277 142 

269-322 256-.^28 272-306 267-320 258-346 129-173 

293 282 290 291 296 141 

265-304 248-289 256-289 253-331 248-3111 138-153 

281 269 277 288 276 145 

270-310 269-306 245-317 276-333 290-306 137-158 

285 287 277 295 299 149 

259-322 241-328 245-347 253-333 243-346 125-173 

284 280 284 287 288 142 

Head length 

279-313 279-290 288-314 290-311 — 202-226 

297 285 301 299 — 211 

284-305 276-302 279-300 278-309 275-313 199-227 

297 288 289 294 291 214 

279-319 255-327 294-339 271-334 273-341 209-226 

306 294 313 300 307 217 

286-323 272-291 289-313 281-316 245-311 200-229 

302 281 299 294 283 211 

306-325 284-311 281-320 298-327 281-336 206-231 

311 297 307 308 306 222 

279-325 255-327 279-339 271-334 245-341 199-2.^1 

302 291 302 299 298 215 



304-351 285-330 292-331 — 

327 301 312 — 

292-330 276-309 270-301 257-307 

311 290 287 286 

31()_37| 303-345 275-318 264-332 

333 324 303 295 

305-361 292-345 28f)-324 249-325 

329 317 308 297 

299-345 298-329 284-317 288-315 

331 311 301 297 

292-371 276-345 27t)-33l 249-332 

328 309 303 294 

Caudai-pedimcle depth 

116-132 121-145 122-144 — 

126 134 131 — 

131-158 119-148 126-153 128-145 

144 1311 137 135 

Il')-I71 123-155 111-182 114-158 

143 133 139 133 

139-165 123-158 131-164 124-154 

148 145 150 140 

138-168 124-166 150-158 135-158 

153 142 152 148 

116-171 119-166 111-182 114-158 

144 137 142 137 

Head depth 

194-223 203-221 198-219 — 

208 213 208 — 

198-222 200-222 199-218 175-213 

208 210 207 200 

191-241 203-233 193-220 192-229 

208 218 208 206 

194-213 202-222 192-221 177-221 

205 212 210 203 

202-221 1^*9-223 210-231 203-230 

216 212 214 216 

191-241 199-233 192-231 175-230 

209 213 210 206 



(Table continued on next two pages.) 



216 CALIFORNIA ACADEMY OF SCIENCES MEMOIRS 

TaBI E 36. CONTINUED. 



Snout length Orbit length 



Sex Males Females Males Females 



S.L., mm. 24-42(-) 42-66 -10-42(-) 42-66(-) 66-114 24-42(-) 42-66 .10-42 (-) 42-66(-) 66-114 



Lake Franklin 

Ruby Valley 7.^-91 72-81 77-87 79-88 — 64-92 5»-15 67-79 .S9-72 — 

81 77 8.1 84 — 78 68 7.1 64 — 

Biillc Valley 73-8."! 68-80 6.'^-77 72-87 70-95 62-81 .59-73 64-81 .'59-72 46-59 

77 74 73 79 79 70 65 70 6.1 52 

Lake Waring 75-89 64-92 65-90 70-104 68-101 66-91 52-75 70-90 49-76 43-66 

81 80 86 S5 89 78 66 78 62 54 

LakeSteptoe 79-92 76-87 79-90 77-93 73-97 65-77 55-68 61-70 51-65 44-59 

85 82 85 85 85 70 61 64 57 50 

lake Spring 78-89 74-87 81-92 79-94 80-96 67-81 61-72 67-90 57-67 51-63 

83 82 86 86 88 74 65 78 62 56 

Total range 73-92 64-92 65-92 70-104 68-103 62-93 52-75 61-90 49-76 43-66 

Grand average 82 79 83 84 86 75 65 73 61 53 

Upper-jaw length Mandible length 



Lake Franklin 

Ruhy Valley 79-91 74-87 K2-88 79-95 — 98-122 94-109 101-116 106-121 — 

85 81 85 86 — 109 lol 110 110 — 

ButlcValley 68-90 65-89 71-86 83-88 78-94 89-114 102-111 104-112 102-111 97-116 

80 81 79 85 88 105 108 107 108 108 

lake Waring 78-94 67-102 79-97 72-97 75-112 1(10 121 87-126 106-132 94-124 91-110 

84 81 88 85 94 111 108 117 110 114 

LakeSteptoe 69-88 74-87 72-85 71-100 75-106 99-119 95-114 97-122 100-122 88-122 

80 79 80 86 87 110 104 107 108 102 

LakcSprinc 81-99 71-85 74-95 77-94 87-109 105-126 99-115 99-122 99-119 99-122 

89 81 85 85 92 114 108 110 109 109 

Total range 68-99 65-102 71-97 72-100 75-112 89-126 87-126 97-112 94-124 88-110 

Grand average 84 82 81 85 91 11(1 107 110 109 109 

Interorbital width Suborbital width 



Lake Franklin 

Ruhy Valley 91-108 89-99 92-107 97-105 — 11-46 41-44 16-47 40-47 — 

97 94 101.. 101 — 40 42 42 44 — 

Bulle Valley 82-100 81-101 86-94 89-101 76-99 12-44 14-42 31-49 l(i-46 36-51 

41 1)0 90 91 88 19 19 18 41 45 

Lake Warmg 91-114 80-112 97-117 85-120 82-111 15-54 12-48 16-51 16-48 18-59 

1114 45 109 101 97 40 40 44 42 48 

Lake Sicptoe 94-107 88-102 91-105 87-104 85-101 19-49 42-49 19-48 40-55 18-55 

1111 45 97 97 96 44 45 41 45 46 

Lake Spring 90-105 85-114 84-120 91-117 94-115 15-46 17-45 17-46 19-52 38-48 

47 100 105 105 106 19 42 41 41 45 

Tolal range 82-114 80-114 84-120 85-120 76-115 32-54 12-49 31-51 16-55 16-59 

Grand averaue 99 95 100,„ 100 97 40 41 42 43 47 



:Tahlc <.i'ni-ludL-d on next pa^f. ) 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



217 



Table 36. continued. 



Depressed dorsal fin 



Caudal-fin length 



Sex 



Males 



Females 



Males 



Females 



S.L.. mm. 24-42(-) 42-66 30-42(-) 42-66(-) 66-114 24-42(-) 42-66 30-42(-) 42-66(-) 66-114 



Lake Franklin 




Ruby Valley 


204-245 
238 


Butte Valley 


233-274 
257 


Lake Waring 


220-263 

245 


Lake Steptoe 


220-244 
230' 


Lake Spring 


228-281 
250 


Total range 


204-281 


Grand average 


243 


Lake Franklin 




Ruby Valley 


166-245 
211 


Butte Valley 


203-236 
219 


Lake Waring 


194-264 


Lake Sieptoe 


164-187 
176' 


Lake Spring 


169-247 
217 


Total range 


164-264 


Grand average 


211 



238-247 207-237 186-216 
242 221 205 

223-264 196-243 185-228 
245 220,, 211 

206-277 195-246 173-227 
234 215 202 

216-271 203-235 194-229 
242t 218 207,., 

240-273 205-245 189-219 
252 218 206,, 

206-277 195-246 173-229 

240v„ 218,,, 205;:. 

Pectoral-fin length 



— 


240-284 


— 


261,,.. 


182-207 


233-282 


194 


259,, 


159-210 


223-276 


1 88:, , 


247,., 


1 70-2 1 6 


216-263 


198 


244,' 


177-213 


226-263 


199,,, 


2457 


159-216 


216-284 


193v-, 


25 3« 



261 247-277 234-253 

261, 261, 242,,, 

225-273 226-247 208-252 

249 239-. 236,, 

209-257 218-253 202-252 

235..- 239^ 228..7 

225-255 214-260 210-279 

240,, 238„ 234,,, 

224-266 212-251 224-241 

240,, 238., 232 

209-273 212-277 202-279 

239,., 243,.-, 233,2 

Pelvic-fin length 



209- 


^249 


-): 


!4 


178- 


-246 


209,,. 


192- 


-231 


-t 


17,., 


203- 


-249 


227 


178- 


-249 


2 


18n 



202-226 157-192 170-181 
213 176 176,,, 

208-242 150-213 151-199 
223 175 178 

188-242 163-187 143-210 
215 178,, 181 

187-232 160-191 167-197 
209 174 182 

199-252 156-190 168-195 
228 172 181 

187-252 150-213 143-210 

218 175,s 180,,, 



— 


138-175 


— 


158 


163-188 


156-185 


177 


170 


147-190 


147-183 


170,, 


167 


144-210 


132-146 


175 


138' 


155-191 


130-194 


179 


166 


144-210 


130-194 


1 74,,, 


160 



152-171 127-152 122-140 — 

163 137 132 — 

155-194 115-164 137-175 123-139 

168 137 155 131 

146-190 119-151 106-169 107-151 

163 136 137 129:,, 

143-188 116-146 130-148 114-148 

167 134 139 133 

157-190 125-145 125-145 121-154 

174 136 138 135 

143-194 115-164 106-175 107-154 

166 136 139 131,0 



^ The lengths of the fins in this series pre low, because the specimens are only 27-.^7 turn. long, and liad presumably not yet developed the 
longer fins characteristic of adults. 



Morphometry. A total of 342 specimens 
were subjected to 1 8 measurements, including 
standard lengtii (tables 36, 37). A total of 98 
were from the basin of pluvial Lake Franklin. 43 
from Collection 2 in Ruby Valley, to represent 
the dwarf population, and 55 from two Collec- 
tions (7, 10) in Butte Valley (all those measured 
from Butte Valley came from below the Lake 
Gale basin ) . From the area of the direct tribu- 
taries to pluvial Lake Waring 128 were utilized, 
from four Collections (16-19), including 60 from 
Collection 17, which contained the grotesque 



oversized females ( fig. 51). From the drainage 
basin of Lake Steptoe 65 specimens were mea- 
sured from Collections 20, 22, 25, and 26. From 
Spring Valley, into which it is held that the species 
was introduced, 51 specimens were utilized, all 
from Collection 32. These scries were selected 
to provide a cro.ss-section of potential variates 
throughout the range of the species (table 37). 
The specimens were further selected to represent 
the smaller males, 24 to 42- mm. in standard 
length (67 specimens); the larger males, 42 to 
66 mm. lone (71 ); the smallest females, 30 to 



;18 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Tari.e 37. MciU'iitil <)/ KclictLis soliliiriiis nlilircd for prapoi tiniKil nicdsnicnicnls. 



Pluvial lake 


Fi 


-anklin 






Waring 








Steptoe 
Steptoe 






Spring 
Spring 

32 




Valley 


Ruby 

2 


Butte 




Goshute 




Total 

Number 


Collection No. 


7 


10 


16 


17 


18 


19 


20 


">") 


25 


26 


28 


Males 






























24-4: - mm. 


19 


1(1 


— 


6 


10 


1 


— 


— 


— 


— 


10 


— 


II 


67 


42-fi6 mm. 


3 


13 


— 


10 


IS 


9 


— 


— 


— 


— 


8 


— 


10 


71 


Females 






























.^(1-42 - mm. 


III 


10 


— 


4 


7 


— 


— 


— 


— 


— 


10 


— 


9 


50 


42-66 - mm. 


1 1 


4 


1 


y 


1(1 


7 


3 


1 


6 


3 


5 


-> 


10 


77 


66-1 14 mm. 


— 


S 


4 


11 


\> 


6 


s 


4 


4 


s 


t 


S 


1 1 


77 


Totals 


43 


51) 


s 


40 


6'J 


23 


5 


5 


10 


5 


35 


10 


51 


342 



42- 111111. long (50); the iiiei.iiiini-si/ed females, 
42 to 66- mm. long ( 77 ) ; an(.l the largest females, 
66 to 114 mm. long (77). The two se.xes and 
the five size groupings were thus well repiesentecl. 

Because the relict dace rather surprisingly 
tends to preserve very soundly and with little 
distortion by opisthotonus or otherwise, and be- 
cause large series were obtained, it has been 
feasible to select, from each Collection iitili/ed. 
specimens well suited for precise measurement. 
Furthermore, the material is highly comi">arable, 
becatise of similar methods and length of preserva- 
tion, and because all the measinements were made 
by one of us ( C.L.H. ). 

The measurements for each of five body parts, 
seven head parts, and four fins were transformed 
into pcrmillage o\ staiulard length, aiul for each 
size group of each collection the ranges and means 
were computed. These ranges and means have 
been tabulated for each of the pluvial lake drain- 
age basins, with the Ruby Valley and Btitte Val- 
ley sections of the Lake Franklin watershed 
separated. Five zoogeographical divisions were 
thus obtained, plus another for the total range 
and grand average for each size-group category 
( table .>6, which omits the head-width propor- 
tions, which are subject to error and would add 
little or nothing to the picture). 

In an effort to determine whether there are im- 
portant differences in morphology between the 
populations from the two pluvial drainage sys- 
tems, or between the pi>pulations within either or 



both systems, individual localities were compared, 
and smaller size groupings were tried. Little 
advantage was realized from either refinement. 
In addition, untransformed measurements were 
plotted against the standard length. For Collec- 
tions 1 6 aiicl 1 7. lines were fitted by eye to indicate 
the changes with increasing size of fish. Then 
the measurements for other Collections were 
plotted against those lines. This was done chiefly 
for Collection 2. the main one from Ruby Valley, 
which seems to be one of the more distinctive, 
and is dwarfed. For predorsal length, head 
length, mandible length, and caudal-|iedimcle 
depth, the proportions for Ruby Valley are low, 
but the higher values roughly match the mean 
line for Collections 16 and 17. The caudal-fin 
measurements are high for the Rtiby Valley lot, 
but the low values appro.ximate the mean for those 
Collections. These differences are perhaps largely 
relatable to the dwarfing of the Ruby Valley fish 
(p. 204) and to the tendency for exuberant de- 
velopment at Johnson Ranch, particularly at Col- 
lection 17 (pp. 2()(S. 214). In any event, the dif- 
ferentiation docs not appear to have reached a 
level indicative of the subspecies distinction. The 
graphs have been retained, but are not herein re- 
|iroduced. 

Somewhal the same degree of difference aiul 
overlap appears when the proportions are com- 
pared by ranges and means, for the data compiled 
by lake basins and valleys (table 36). The dif- 
ferences appear to be about as great within as 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



219 



Table 38. Size frequemies by sex and mcitiirily in five representalive Collections of Relictus solitarius. 



Standard lengths by two units of 0.5-cm. size classes 



Pluvial lake system 
Valley (Coll. No.) 

(Date) Sex' 



1.5 



2.0 

+ 



3.0 

+ 
3.5 



4.0 

+ 
4.5 



5.0 

+ 
5.5 



6.0 

+ 
6.5 



7.0 

+ 
7.5 



8.0 

+ 
8.5 



9.0 

+ 
9.5 



10.0 11.0 

+ + 

10.5 11.5 



Lake Franklin 

Ruby ( : » 
(Sept. 12) 



Butte (7)- 
(June 27) 

Lake Waring (17) 
(June 22-23) 



Lake Steptoe ( 25 ) 
(Aug. 23) 



Lake Spring (32 ) ' 
(Julv A) 



Yg. i + 9 
Imm. i 
Imni. 9 
Ad. i 
Ad. 9 

Ad. S 
Ad. 9 

Imm. (J 
Imm. 9 
Ad. c5 
Ad. 9 

Yg. ,^ + 9 
Imm. i 
Imm, 9 
Ad. i 
Ad. 9 

Imm. i 
Imm. 9 
Ad. c? 
Ad. 9 



4 27 

— 17 

— 32 



10 

2 

1 



71 
36 



4 
65 
23 

2") 
4 

4 

13 

119 

62 



13 

■> -) 

48 
26 



30 
99 

33 



192 
147 



15 
14 



4 
10 



87 



21 



10 



1 
46 
17 



5 
130 



39 
32 



5 

75 



^ Unless sex and maturity was obvious by inspection of sexually dimorphic characters, gonads were examined. The distinction between immature 
and maturing is somewhat tenuous and probably not wholly consistent. 

-This collection also included an apparent intersex (labelled no. 16). 6.^ mm. long. 
■'This collection also included an apparent intersex, 53 mm. long. 



between the two main pluvial drainage complexes. 
In general, the deviation of any mean value for 
any dimension from the grand average for tiic 
species is relatively slight, in comparison with 
the wide range for the pertinent sex and size 
group. 

Sexual dimorphlsm and nuptial charac- 
ters. One of the remarkable features of RcUcuis 
solitarius is the marked discrepancy in the average 
and the maximum size of the sexes. This has been 
evident from casual examination of the Collec- 
tions in general, and is illustrated by the sizes 
of the specimens subjected to proportional mea- 
surements (tables 36, 37). The discrepancy in 
size is documented by an analysis, according to 
sex and maturity, of all specimens in five repre- 
sentative Collections (tables 38, 39): Collection 
2 for the Ruby Valley and Collection 7 for the 
Butte Valley divisions of the drainage basin of 
pluvial Lake Franklin, Collection 17 for the 



Goshute Valley and Collection 25 for the Step- 
toe Valley sections of the drainage basin of Lake 
Waring, respectively, and Collection 32 for Spring 
Valley. Among these five Collections, the values 
for the males, expressed in percentage of the value 
for the females, ranges for the mean size from 90 
to 100 for the immature fish and from 68 to 86 
for the mature ones. The corresponding values 
for the maximum size were little altered for the 
immature fish (86 to 100), but diverged notably 
for the mature ones, from 49 to 76. The lowest 
value is for Collection 17. in which the sex ratio 
is equal; the highest value is for the dwarfed Col- 
lection 2 (table 39). The data for the .same five 
Collections tallied by units of two 0.5-mm. size 
classes (table 38) clearly confirm the larger size 
of the females. In Collection 2, among 232 ma- 
turing fish, the largest male measures only 44 
mm. and the largest female only 58 mm. In the 
four Collections of less dwarfed fish, among which 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Tahi K ?>'■>. Niinihcr and size, hy \c\\ ami niatiirily. of ■ipcclniciis in five representative Collections of Rclictus 
solitarius.' 



Standard length, mm." 



Pluviid lake system 
Vallcv (Coll. no.) 
( Date ) 



Unse.xed very 
young 

Min.-max. 
(means) 



Immature 



Adults 



Min.-max. 
(means) 



Min.-max. 
(means) 



Mean Max. 

ShS/ SL c? / 

Mean Max. 

SL 9 SL 9 

X ino X 100 



Sex ratio, 
.V6 :100 9 



Imm. Adult Imm. Adult Imm. Adult 



Lake Franklin 

Ruby (2) 
(Sept. 12) 

Biitlc ( 7 ) 
(.Inne 27) 



Male 

female 
Male 

Female 

Mall 



I.S-22( |y.4 .) 



^- I 



Lake Waring (17) ■ 

(June 22-2-'') I Female 



Lake Steptoc (25) 
( Auu. 2.M 



Lake Spring (.^2) 
(July 6) 



Male I 

Female J 
Male ] 

Female J 



L^-I7( I.VO 



2.^-28(25.5,,,) 
23-2S(25.6.;0 



17-.^8(29..V) 

l5-4()(.^2.6,.,) 
17-36(21.9:;) 

17-42(2.3.6,.) 
25-.32(29.3,,) 

21-37(29.6,,-,) 



24-44(35.7,,.,) 

30-58(41.7,:,) 
33-66(42.8,,) 

33-99(62.5,,.) 
29-56(39.9,,,:,) 

30-114(46.9,,,) 
34-51(42.8,-.,:) 
38-80(54.4,.,) 
28-53(37.8,,,) 

29-86(53. 6,„,) 



100 86 100 76 53 76 



68 



67 



79 86 64 173 



99 71 



86 



80 



90 85 95 49 37 99 



29 



50 44 



' Frequencies by two units of 0.5-cm. class are entered in table 38; subscript figures denote number of specimens. 
■' Means were computed from data grouped by 0.5-cm. classes. 



1,37U specimens were all iiieasmed. the longest 
males are only 31 to 66 mm. long, the longest 
females 80 to 114 mm. The longest female in 
each of the.se four series is 1 .5 to 2.0 times as long 
as the longest male. The longest females would 
be about 3.5 to 8.0 times as heavy as the longest 
males, if the usual approximate cube relation 
holds. The lesser size of the males seems to hold 
in all other Collections of the species containing 
adult fish. The very restricted habitats and the 
thoroughness of collecting rule out the alterna- 
tive suggestion that the older and larger males 
were unavailable. 

The occasional tKcurrence of apparently over- 
sized and perhaps overage females (fig. 51 ), par- 
ticularly at Collection 17, is discussed under Size 
(pp. 208-209 and fig. 50). 

A comparison of the proportional measure- 
ments for the smaller and larger males, and for 
the smallest, medium, and largest females, by 
geographical areas (table 36), gives some indica- 
tion of some sexual differences in chances with 



growth in the relative size of the body and head 
parts and of the fins, as follows: 

I'rcdorsal Icoglli: appro\inialcl\ isometric in 
males, increasing in females, mtirkcdly so in 
oversized fish (p. 215). 

Anal origin to caudal base: increasing in males, 
except lor the dwtirl population ol Ruby 
Valley: decreasing in leiiiales (presumably by 
reason of an increase in length of abdomen — 
a freqtient occurrence in females). 

Hody depth, caudtil-pedimcle depth, and, con- 
trary to a general trend in fishes, head length: 
all fluctuate without apparent marked trend. 

Orbit length: decreasing in both sexes, as almost 
alwa\s in fishes. 

Upper-jaw length: about isometric in males, 
probably slightly increasing in femtiles. 

Mandible: probtibly decreasing slightly in each 
sex. 

hiterorbittil width: decreasing slightly in each 
sex. 

Suborbital width: about isometric in males, in- 
creasing in femtdes. 

Fins: all decreasing in both sexes. 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



221 



Table 40. Se.xiitil diiiuirphisni in predorsal and fin leni^tlis in Rclictus solitiirius, expressed <is excess for males 
over females in mean values in permilla^e of standard length.^ 



Pluvial lake system 
Valley 


Predorsal 
A B 


length 
C 


Dorsal fin 


Caudal fin 


Pectoral fin 


Pelvic fin 


A 


B 


C 


A 


B 


C 


A 


B 


C 


A 


B C 


Lake Franklin 






























Ruby 


4 


_I3 


— 


17 


37 


— 


I) 


19 


— 


35 


37 


— 


21 


31 — 


Butte 


~1 


-19 


-27 


37 


34 


51 


20 


13 


15 


44 


45 


46 


33 


13 37 


Lake Waring 


-4 


-17 


-28 


30 


32 


46 


8 


7 


30 


44 


34 


45 


31 


26 34 


Lake Steptoe 


-10 


-13 


-IS 


12= 


35 


44 


6" 


6 


23 


-)- 


27 


34 


4-- 


28 34 


Lake Spring 


_-> 


-19 


-29 


32 


46 


53 


7 


8 


13 


45 


47 


49 


30 


36 39 


Total 


-3 


-17 


-23 


25 


35 


47 


10 


6 


21 


36 


38 


44 


24 


27 35 



' Data from tables 36. 37. which indicate number of specimens measured of each size group, two for males and three for females. In column A and 
B males of the smaller and larger size group are compared, respectively with females of the same size groups; in column C the larger males are com- 
pared with the largest size group of females (larger than any males I . 

-Data are aberrant because the fish in the sinall-size category arc only 27-37 mm. long, and had presumably not yet developed much se.xual 
dimorphism in the length of the fins. 



Sexual dimorphism in size of all fins is strongly 
marked in this species. The greater length of the 
fins in the males (table 40). e.xpressed as excess 
in males of the mean value in permillage of stan- 
dard length, is much greater for the dorsal fin 
than in cither Rliinichthys osculiis (table 14) or 
Gila hicolor ( table 28 ) samples — the excess 
values are relatively consistent, closely approxi- 
mating those for the pectoral and pelvic fins. 
The excess values fluctuate from slight to moder- 
ate for the caudal fin. as they do for the two other 
species. The excess values for the pectoral and 
pelvic fins are somewhat more consistent than for 
the other species. In grand average, the pectoral 
is about 4 percent, the pelvic about 3 percent, of 
the standard length longer in the male than in the 
female. Such rather extreme .sexual dimorphism 
may be regarded as an example of exuberance, 
such as we mentioned for certain races of Liicania 
parvci (Hubbs and Miller. 1965, p. 33). 

In the males, the fins are not only elongated 
but are also broadened, and the rays are thick- 
ened, especially along the front or outer edge. 
Thus, the sexes are notably .sexually dimorphic. 

In several respects, the breeding males tend to 
develop the adult coloration at a smaller size than 
the females and carry the development farther. 
This was noted, for example, in the attainment of 



dense puncticulation on the upper and lower lips 
and on the intergular region. 

The characteristics of the nuptial tubercles, as 
briefly indicated in the generic diagnosis (p. 182). 
furnish some of the more distinctive features of 
the relict dace. The following description was 
drawn from males in Collections 17 and 25. but 
at least incipient development of the same type 
was noted for several other Collections, and it is 
assumed that the pattern is species-specific. On 
the head, the largest of the distinctively straight, 
cone-shaped tubercles line the infraorbital branch 
of the lateral line uniserially. The next most 
prominent tubercles line the suborbital margin, 
also uniserially. In addition, very small cones 
are .scattered over the lower-cheek and mandibu- 
lar regions, and. in .some males, a few develop 
just behind and abiive the eye. 

On the pectoral fin rather large but definitely 
caducous and very distinctive erect cones extend 
uni.serially along the upper edge of the topmost 
(outermost) ray. curving to an inner position on 
the ray posteriorly (in Rliiniclulrys oscuhis. the 
much thickened uppermost ray is devoid of 
tubercles; in Gila hicolor. small, firmly attached, 
curved tubercles line the first ray uniserially). In 
addition, similar but usually smaller organs ex- 
lend along the next one to four rays, leaving the 



Ill 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Tahi E 41. Number of fin rays in (crlciin Collections of Rclictus solitarius //; diffcrenl drainai^e basins. 



Pluvial hike system 
Valley (Coll. no.) 



Dorsal Rays' 



Anal Rays' 



9 10 No. Mean 



8 No. Mean 



Lake Franklin 
Ruby (I, .^. .";. 6) 

Butte 1'^' 
I (1 I ) 

Lake Waring (17) 

Lake Steptoe (26. 27) 

Lake Spring (32 ) 

Total 



26 — 

IX — 

15 — 

20 — 

24 1 

19 — 

122 1 



31 


7.93 


20 


7.90 


15 


8.00 


20 


8.00 


26 


8.00 


20 


7.95 



132 



7.96 



1 


30 


— 


31 


6.93 


1 


19 


— 


20 


6.95 


3 


12 


— 


15 


6.80 


- 


20 


— 


20 


7.00 


1 


23 


T 


26 


7.04 


1 


19 


— 


20 


6.95 


7 


123 


T 


132 


6.96 



■■■luvial lake system 
Valley (Coll. no.) 



Caudal Rays- 



Pelvic Rays' 



16 17 18 19 20 21 No. Mean 



9 No. Mean 



Lake Franklin 

Ruhv (2) 

Buttc \']\ 
I ( I I ) 

Lake Waring (17) 
Lake Steptoe ||;^| 
Lake Spring ( 32) 
Total 



- 20 — — 21 18.86 

1 14 2 1 18 19.17 

1 12 — — 13 18.92 

3 13 3 — 20 18.90 

1 14 2 1 18 19.17 

2 9 1 — 14 18.64 
8 82 8 2 104 18.95 



7 


39 


4 


50 


7.94 


4 


34 


") 


40 


7.95 


1 


28 


1 


30 


8.00 


— 


29 


11 


40 


8.27 


4 


30 


6 


40 


8.05 


8 


177 


20 


228 


7.93 


S 


32 


3 


40 


7.95 



44 369 47 468 7.98 



Pluvial lake system 
Valley (Coll. no.) 



Pectoral Rays' 



9 10 11 12 13 M 15 16 No. Mean 



Lake Franklin 

Ruby (2) 

Butte 1<^' 
|( 11 ) 

Lake Waring (17) 

Lake Steptoe (26) 

Lake Spring (32) 

Total 



1 


14 


17 


5 


3 


40 


13.87 


— 


8 


")-) 


s 


5 


40 


14.17 


-) 


10 


9 


7 


-> 


30 


13.90 


3 


16 


20 


1 


— 


40 


13.48 


") 


9 


12 


15 


1 


40 


13.97 


4 


16 


10 


4 


6 


40 


13.80 


12 


73 


90 


37 


17 


230 


13.87 



' Last 2 elements counted as 1 ray. 
-■ Principal rays (branched rays +2). 
■ Both sides citunled. 



proximal and distal parts of the lays unarmed. 
The files remain uniserial throughout the ray, not 
branching, as they diagnostically do in Rliinich- 
thys and (Jila. In high males, similar but weaker 
tubercles occur along each of the outer one to 
several pelvic rays and along each of the anterior- 
most one to a few anal rays. No nuptial tubercles 
ever develop on the body. 

In nuptial males the fin rays are considerably 
thickened, especially along the anterior (or upper 
or outer) edge of the fin. 

Fin rays (table 41 ). As in Rlilnichiliys and 
Gild hicolor, the dorsal rays rather seldom vary 



from 8; when they do, the number is more often 7 
than y or 10. As in Rhinichlliys osciiliis, the anal 
rays ( 6-X ) seldom deviate from 7: the usual num- 
ber in Gila hicolor is either 7 or 8. The caudal 
rays vary widely, from 16 to 21, about as in the 
Rliiiiiclilhys osculns and Gila bicolor samples, but 
in this species the deviates from the family norm 
of 19 are about as frequently up as down. The 
pectoral rays vary in number from 12 to 16, with 
a single count of 9, and with a grand-total mean 
of 1.^.87, lower than for any of the samples of 
Gila hicolor treated; about equal to most means 
for the samples of Rhiniclilhys osciiliis rohiisliis. 



VOL. VII HUBBS, MILLER. & HUBBS— GREAT BASIN RELICT FISHES 



223 



Table 42. Niimher of vertebrae in certain Collec- 
tions of Relictus solitarius in different drainage basins. 



Pluvial lake system 
Valley (Coll.no.) 






Frequencies' 




35 


36 


37 


38 


39 


No, 


Mean 


Lake Franklin 
















Ruby (2) 
Butte -1<^' 

|(in 


-> 
1 


5 
14 


6 

3 
4 


1 
5 


I 


13 
19 
12 


36.85 
36.05 

37.08 


Lake Gale 


— 


4 
3 


8 
6 


1 
1 


— 


13 
10 


36.77 
36.80 


Lake Waring 


-» 


8 

7 


6 


1 


— 


14 
12 


36.43 
36.17 


Lake Steptoe 
steptoe |;-| 


1 

1 


6 
9 


8 
4 


1 


— 


15 
15 


36.47 
36.33 


Lake Spring 
Spring (33) 


— 


10 


4 


T 


— 


16 


36.50 


Total 


7 


68 


51 


12 


1 


139 


36.51 



1 Including hypural complex as 1 vertebra and including the tour 
comprising the Weberian apparatus. 



but higher than the means for the isolated sub- 
species. The pelvic rays range from 7 to 9 and 
usually number 8, as in the isolated spring popu- 
lations treated of GiUi bicohn- ( modally 9 in G. 
b. obesa) and as in the Humboldt Valley popula- 
tions of Rhinichthys osctiliis robiistiis ( usually re- 
duced to 7 in isolated spring samples of R. 
osculiis). In summary, the ray numbers show 
some evidence toward reduction in this spring- 
inhabiting fish. No marked regional variation is 
noted. Ray counts in the holotype are D 7. A 6, 
P, 14—13. Pl. 8—8. C 19. 

Vertebrae. The vertebrae (table 42) num- 
ber 35-39, usually 36 (36 in holotype) and/or 
37 (37 or 38 in one Collection), much as in the 
forms of Rhinichthys osciilits treated (35-40. usu- 
ally 36-39), somewhat less than in the subspecies 
of Gila bicolor handled (37-42, usually 38-4(1). 
Of the ten representative Collections counted, 
only one of two from the section of Butte Valley 
that drained directly into Lake Franklin shows 
a noteworthy divergence from the others. 

Scale rows. Numbers of scales in the five 
rows counted ( table 43 ) usually run somewhat 



higher than in the forms studied of Rhinichthys 
osculiis, but lower than in those of Gila bicolor. 
There is considerable local variation, but overlaps 
in counts are generally wide and some of the 
counts are not very precise for reasons cited (p. 
91 ). In general, counts for the drainage basin 
of pluvial Lake Franklin, especially those for the 
apparently dwarf form of Ruby Valley, tend to 
run lower than those for the basin of Lake 
Waring, but the differences are not sharp enough 
to warrant subspecific separation. The circum- 
stance that the sample from Spring Valley agrees 
in scale count best with the samples from the Butte 
Valley .section of the ancient drainage basin of 
Lake Franklin, is cited as an indication that the 
.source of the assumed introduction into Spring 
Valley was one of the springs of Butte Valley (p. 
235). In the holotype, the scale row counts (all 
approximate, by reason of irregularities) are: in 
lateral line, 60; above lateral line, 13; below 
lateral line (to anal origin), 12; lateral line to 
pelvic insertion. 9; predorsal, 35; around body, 
26 above, 31 below, 59 total. 

Gill-rakers. The rakers (table 45 ) vary from 
7 to 12 and usually number 8 to 11. With some 
overlap, they are more numerous than in Rhinich- 
thys osculiis. but fewer than in Gila bicolor. They 
arc also larger and better developed than in R. 
osculiis. but smaller and weaker than in G. bi- 
color. The counts average slightly lower for the 
Butte Valley .section of the drainage basin of Lake 
Franklin and for Spring Valley than for other 
parts of the range, as do the scale counts, further 
suggesting that the Spring Valley population may 
have been introduced from Butte Valley. 

Pharyngeal arch and teeth. These struc- 
tures are discussed above under the heading of 
the genus. The data (table 33) do not suggest 
any regional differentiation in tooth number. 

Karyotype. Treated under the genus heading 
(p. 193). 

Sexual differences in numbers and bio- 
mass. Among the five large and representative 
Collections for which data on population structure 
were accumulated (tables 38. 39). the sex ratio 



224 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



TabI-E 43. Niiiiihcr of scdlc laivs unit scii.sury pores in i ciniiii CCHectinns of Rclictiis solitarilis //( different drnin- 
cific biisins. 



Pluvial lake system 
Valley 



Number of scales in each row- 



Coll. (No.)' 



Lateral- 
line rows 



Predorsal 
rows 



D-A Around 

origins body 



Around 
peduncle 



Number of pores' 

Lateral Supra- 
line temporal 



Lake Franklin 

Ruhv 



Butte 
Lake Gale 

Lake Waring 

Lake Stcptoe 

Lake Spring 
Total 



2 (5) 

7 (5) 

II (10) 

14 (10) 

]h (10) 

17 ( .<! ) 

2.*; (10) 

26 ( .M 

32 (5) 



5 I -60 
55.2 

50-57 
54.6 

5 1 -65 
5S.5 

54-65 
60.4 

57-66 
62.2 

58-66' 
60.3 

55-66 
60.8 

58-70 
61.8 

50-55 
52.6 

50-70 
54.1 



32-34 
32.6 

30-33 
31.4 

27-36 
32.5 

32-36 
34.3 



3 1-34 

33.4 



30- 



30-33 
31.2 

27-34 
32.8 



20-23 
21.6 

21-23 
22.4 

21-26 
23.3 

22-24 
22.8 



22-26 
24.8 



25-28 
26.2 

21-25 

22.4 

20-28 
23.3 



52-60 
54.8 

55-58 
55.8 

55-64 
60.2 

54-64 
58.0 



60-66 
62.6 



59-61 

59.8 

55-61 
58.6 

52-66 

58.7 



28-30 
29.2 

30-31 
30.2 

28-30 
29.6 

30-32 
30.8 



30-34 
32.0 



32-34 
33.0 

29-32 
30.4 

28-34 
30.6 



3-17 
11.8 

13-26 

18.4 

6-21 

12.9 

11-29 
17.9 



11-26 
16.7 



3-25 
17.0 

14-25 
18.2 

3-24 
16.0 



2-4 
3.2 

2-4 
3.0 

2-5 
3.4 



3.2 



0-5 
3.0 



3-4 
3.4 

2-4 
3.1 

1-5 



' Collection number, and. in parcntlieses. number of specimens, except as indicated by footnote. 

- For each row. tlie minimum and maximum counts, and the means. 

■'• Both pore counts were made on botli sides, doubling the number of counts. 

' 15 specimens. 



vaiiecl ticiiicndously and unaccountably: t'rcini 37 
to 173 males per 100 females for the immature 
fish and from 2^) to 99 for the mature fish. In 
view of the greatly restricted habitats and the 
thorough collecting, it seems improbable that dif- 
ferential schooling or different habitat directly 



caused the variation. Differential mortality or 
some chromosomal aberration would seem to pro- 
vide a more plausible explanation. Rather small 
samples may have resulted in some of the fluctua- 
tion for the immature fish, but the samples seem 
adeciiiate foi- the mature specimens. Oddly, in- 



Tabi.E 44. C('iinl\ of nunulihiilur pores in po/uilii- 
lions (if RclictLis solitariiis in lerfnin Inisins in IMewulci. 



Tabi i; 45. Number of i^ill-rnkers in certnin Collee- 
lions of Rclictiis solitarilis //; Jifferenr tlr(nnci:^e basins. 



Pluvial lake system 
Valley (Coll. no.) 3 


4 


Mandibi 
5 6 


liar 

7 


Pores 
8 No. 


Mean 


Pluvial lake system - 
Valley (Coll. no.) 








Gill 


-rakers 






7 


8 


9 


10 


11 


12 


No. 


Mean 


Lake Franklin 














Lake Franklin 


















Ruby (2) — 
Butte (7) — 


5 
4 


24 
21 


9 


1 
-> 


1 40 

— 41) 


5.23 
5.33 


Ruby (2) 

Btttte 1'^; 
|(1 1 ) 


1 


4 

1 


7 
12 
11 


12 
3 


1 
1 


— 


20 
20 
15 


9.70 
8.90 
9.13 


Lake Waring (17) — 


4 


15 


12 


6 


1 38 


5.61 


Lake Waring (17) 
Lake Steptoe (26) 


— 


5 


6 

4 


7 
9 


-> 

6 


1 


20 
20 


9.30 
10.20 


Lake Steploe (26) 1 


6 


28 


5 


— 


— 40 


4.93 


Lake Spring (32) 


— 


7 


10 


3 


— 


— 


20 


8.80 


Total 1 


19 


88 


39 


9 


2 158 


5.27 


Total 


t 


17 


50 


36 


10 


1 


115 


9.35 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



225 



deed. Collection 1 7 with the lowest proportion 
(37) of males among the immature fish (un- 
fortunately a small sample ) shows a balanced 
representation of sexes among the adults; whereas 
Collection 25, with 1.7 males to 1.0 females 
among the immature fish, had the most un- 
balanced sex ratio among the adults ( 3.4 females 
to 1.0 males). 

In as much as the females average larger than 
the males, and ordinarily outnumber them, the 
contrast in the total biomass of the sexes must be 
large. 

Abundance and life history. Numerous 
field observations and local testimony indicate 
that Relictiis solitaritis is a very prolific fish, with 
a rather long breeding season, extending at least 
from late June to mid September (the entire 
period of collecting). The spawning period is 
roughly approximated by the frequent collection 
or observation of the very young at various dates, 
as indicated by the minimum size reported above 
in the account of the Collections (pp. 199-207). 
For example, extremely small fish seen at Collec- 
tion 4 on September 1 3 suggested that spawning 
was still in progress. Inclusion of nuptial males 
and of gravid females at various times in a num- 
ber of Collections confirmed the postulate of an 
extended spawning period. 

The prolonged spawning may be related to the 
moderately warm and presumably relatively con- 
stant temperatures of the type of spring waters 
inhabited by this species ( see below ) . 

Great abundance was shown by the very large 
numbers collected — at times in single hauls of a 
small seine (as is also indicated in the descrip- 
tions of the Collections). Former swarming where 
the dace have become depleted or extirpated by 
agriculture, fish-stocking, and industry is indi- 
cated in the section on Material Examined and 
Population Status: in the general remarks on 
Populations of Ruby Valley; following the ac- 
count of Collections 10 and 19; in the discussion 
(pp. 204-207) of Populations of Steptoe Valley, 
especially in the local testimony ( p. 205 ) "that 
wagon loads could be gathered when the holes 
went dry in summer." 



That the relict dace deposits its eggs on plants 
is suggested by its close association with sub- 
merged vegetation growing on soft, more or less 
anaerobic bottom, in regions where any substrate 
other than the plants is unsuitable. 

The size-frequency distribution at representa- 
tive Collections (table 38), confirmed by inspec- 
tion of other series, indicates, with the dates of 
collection in mind, that both sexes spawn first as 
yearlings, but that the smallest yearlings tend to 
reproduce late in the second year of life. The 
rather compact size distribution of the mature 
males suggests that few males breed at an older 
age. That the females grow so much larger than 
the males is an indication that many of them 
breed when two or more years old. No attempt 
was undertaken to determine age by a study of 
annuli on scales, otoliths, or bones. 

Habits. The relict dace is typically a midwater 
swimmer, in agreement with its body form (p. 
212). It is seldom seen either at the very surface 
or resting on the bottom. 

It is one of the most secretive of minnows — 
more so, we think, than any other western species. 
On even slight disturbance it tends to dive 
promptly into the soft bottom or into the thick 
patches of Churn or other submerged plants (usual 
features of its habitat; see below). Once there, it 
may remain out of sight for a considerable period. 
These habits were particularly striking when first 
ob.served on September I 2. 1 934, evoking a field 
note, for the springs and open marshes of Ruby 
Lake. Here, the secretiveness may have been ac- 
centuated as an escape behavior, in the pressure 
from the abundant waterfowl. 

That the relict dace may become especially re- 
tiring during cold periods is suggested by observa- 
tions on September 13-14, 1934, when water 
froze in our camp overnight. No minnows were 
seined or seen on a careful observation of the 
spring-fed waters from about 2 km. south of 
Currie to the source, in a pond about one-half 
acre in size, or in two tributary groups of mound 
springs just to the west. The pond at the base of 
lava hills had the typical features of Relictiis 
habitats: heavy growth of Clnini and Nasturtium. 



226 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



algae. Polainogcloii. cf. P. pccliiuitus, cane, etc.. 
with open spaces. That the species eluded ob- 
servation during that cold period is indicated by 
our later seining of 201 specimens in these waters 
(Collection 19). and by local testimony on their 
presence (p. 219). 

The relict dace seems to be a hardy fish. A 
rancher who has used it said that it serves well as 
live bait in sport fishing and that it lives long when 
thrown out on the bank. None died during trans- 
port by car to Michigan. 

Habitat. Our field exploration cimfirmcd the 
local testimony that this minnow occurs through- 
out Butte Valley, ""without variation," in all 
"warm" springs (obviously meaning those that 
are not hot, nor definitely cold, and do not freeze 
in the winter). We found this to be generally 
true throughout the range of the species. It seems 
to be absent in all canyon streams. When the 
species was common in spring waters in Ruby 
Valley, in 1934, we found none in Cave Creek, 
an icy-cold stream that discharged from a large 
cave (mentioned by Wheeler, 1889, p. 26). nor 
in very cold springs issuing from the same lime- 
stone formation nearby — though these waters are 
in contact with those in which the relict dace 
flourished. 

In the spring-fed waters where it cKcurs. it 
tends to concentrate in quiet pools, particularly 
those that are well vegetated, and where the banks 
are undercut. 

The temperature of the water was noted at the 
time of collecting, generally in the hot season, as 
'"cool" or "moderate," or, when measured ( in 
Fahrenheit scale, transposed to nearest Celsius 
degree), as 9 . 12 . and 14 (once each); 15 
and 16 ( twice each ); I 8 ( four times ): 19 and 
20 (once each). 21 ( four times), and 22 , 2."* , 
24 , and 25 ( once each ) . 

The vegetation was recorded as "little" only 
thrice; ""some" or "moderate" (twice); "much." 
"dense." ""generally dense."" or ""rather thick" ( 1.^ 
times), and ""choked" or ""generally choked" (5 
times). The plants were identified roughly as 
Ndslurliiim (water cress). Poldnioi^cton. cf. P. 



pcctiiuiliis. broad-leaved species of Potcnnogelon, 
CInini. L/lriciil<iil(L rush, bur-reed, grass, alga, 
and moss. 

As noted under Habits, above, the relict dace 
on even slight disturbance dives into the thick 
vegetation, and remains hidden for a consider- 
able time. 

Utilization. The ranch operator at Stratton 
Ranch in Butte Valley testified on June 2. 1942, 
that he had taken this minnow from there to Lake 
Mead, where they served satisfactorily as live bait 
for gamefish. It was further stated that it has 
been used as "scent" in trapping coyotes. 
Ranchers said also that it has been eaten, "as 
sardines." 

Another possible utilization of the relict dace 
came to our attention on June 6, 1942, when the 
owner of Morgan Ranch Spring in Rush Valley, 
Tooele County, Utah, who was developing a re- 
sort there, told us that in building up a stock of 
bullfrogs he had utilized a fish that we assume to 
be this species. He had heard that minnows .serve 
as food for bullfrogs and that minnows are com- 
mon at the McCiill Ranch in Steptoe Valley, so he 
went there, three years previously, and brought 
over three barrels of "small blue minnows,"" 
avoiding the ""goldfish." These were almost cer- 
tainly representatives of Rclicttis soUhirius (see 
Collection 25. p. 205). However, hundreds of 
Utah chubs {(Jila ainiria). either native or also 
introduced, were the only fish we saw in his 
spring, and it appeared doubtful that any relict 
dace had survived. The water temperature, about 
29.5 C., may well have been too hot for Rclictits 
( sec above ) . 

The introduction of the relict dace into Spring 
Valley, presumably at Stone House (pp. 23.^- 
235) may have been for mosquito control, or 
perhaps as a curiosity. 

Derivation of namf.. The Latin term soli- 
Uiriiis is defined as '"alone, by itself, lonely, soli- 
tary." It refers to the evidence that Relictiis 
solituriKs is the lone native inhabitant of any of 
the four Pleistocene lake basins in which it nat- 
urally occurs. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



227 



GENUS CRENICHTHYS HUBBS 

Creuichthys Hubbs. 1932, pp. 1-4, fig. 1 (diagnosis; 
comparisons and relationships; type, C. nevadac) . 
Brues, 1932, p. 280 (distinct genus; relationships). 
Sumner and Sargent, 1940, pp. 46-54 (C. haileyi 
recognized by Hubbs; physiology and hot-spring 
habitat). Hubbs, 1941a, p. 66 ( mention of discovery ). 
Hubbs and Miller, 1941, p. 1 (distribution; C. 
baileyi included in genus). Sumner and Lanham, 
1942. pp. 313-327, figs. 1-4 (physiology and hot- 
spring habitat). Hubbs and Miller, 1948b, pp. 90- 
91, fig. 24 (distribution; species). Miller, 1948, pp. 
99-100 (relationships; distribution). Kopec. 1949, 
pp. 56-61 (history of genus and species; biology of 
C. baileyi). La Rivers. 1952. p. 90 (characters in 
key; compared with Eiupcirichthys: "Springfish" ) . 
Eddy, 1957. p. 162 (characters in key; desert streams). 
Moore, 1957. pp. 150-151 (comparisons, in key; 
teeth; species; range). Miller, 1958, pp. 206-207. 
figs. 15, 16 (two species; distribution as isolated 
relict). Hubbs and Drewry, 1962, pp. 107-110 
(artificial hybridization regarding relationships). La 
Rivers, 1962, pp. 21, 31, 87, 109, 512-516, 681-686 
(range; environment; conservation; review of litera- 
ture). Uyeno and Miller, 1962, pp. 520-532, figs. 
2B, 38, 58, 7 (relationships; osteology and dentition; 
distribution). Bradley and Deacon, 1965, pp. 55 
and App. IL p. 2 ( habitat and community of both 
species). Hubbs and Hettler, 1965, pp. 245-248 
(tolerance to environment). Hubbs, Baird, and 
Gerald, 1967, pp. 104-115, figs. 1-3 (activity cycles 
related to diurnal fluctuations in dissolved oxygen). 
Moore, 1968, pp. 109-110 (comparisons, in key; 
diagnosis; species). Hubbs, 1970, pp. 295-296 (arti- 
ficial hybridization regarding relationships). 

Additional papers bearing on the responses of Cre- 
nichlhys to its environment, and on the threat to this 
native genus resulting from the introduction of exotic 
fishes, but dealing with Creuichthys baileyi, are: Deacon, 
Hubbs and Zahuranec, 1964; Hubbs and Deacon, 1965; 
Wilson. Deacon, and Bradley, 1966; Deacon and Wilson, 
1966 and 1967; Cole, 1968, p. 477; Minckley and Dea- 
con, 1968. 

We have already referred to thi.s eyprinodont 
genus as one of the two types of fish that inhabit 
the warm springs of Railroad Valley, in which 
pluvial Lake Railroad accumulated, and the long 
chain of warm springs in the remnant course of 
pluvial White River (p. 35), just to the east. In 
addition, we have referred to it, further (p. 71 ). 



as one of the inhabitants of the north-central 
Great Basin. 

We have verified the specific distinctness of the 
two forms of Creuichthys: C. nevadae of Rail- 
road Valley and C. baileyi of the White River 
remnants. 

The status and relationships of the relict 
cyprinodontid genera Creuichthys and Empetrich- 
thys of the Great Basin have been discussed over 
the past 40 years, with reference to earlier supposi- 
tions, by Hubbs (1932. pp. 1-4), Hubbs and 
Miller (1941. pp. i-2). Miller (1948. pp. 99- 
100). Uyeno and Miller (1962), and Hubbs and 
Drewry (1962). On the basis of hybridizing 
potential. Hubbs ( 1970, pp. 293-296) suggested 
that Creuichthys and the alien, western species 
of Fiindiihis. F. parvipinnis Girard, are closely 
related. This view indeed seems to make .sense, 
in view of the zoogeographical and paleontologi- 
cal data. Without entering here into a renewed 
study of Creniclitliys and its relationships, we have 
merely brought together the preceding annotated 
synonymy of the genus. 

Railroad Valley Springfish 

Creuichthys nevadae Hubbs. 

Creuichthys uevadue. Hubbs, 1932, pp. 1-6, pi. 1 
(original description of genus and species; isolated 
warm spring at Duckwater. Nye County, Nevada). 
Brues. 1932. p. 280 (type habitat). Sumner and 
Sargent. 1940. p. 46 (mention). Hubbs, 1941a. pp. 
66-67, fig. 5( collection of types; abundance in hot 
springs of Railroad Valley). Hubbs and Miller, 
1941, pp. 1-2 (type of Creuichthys: in hot springs 
of Railroad Valley, replacing C. baileyi: compared 
with C. baileyi): 1948b, pp. 90-91, 93. fig. 24 (hot 
springs in Railroad and Duckwater valleys). Kopec, 
1949. p. 56 (discovery of species). La Rivers, 1952, 
p. 91 (characters in key; compared with C. baileyi: 
Railroad Valley system, west of White River; "Rail- 
road Valley Springfish"). La Rivers and Trelease. 
1952. p. 118 ("Railroad Valley Springfish"; counter- 
part of C. haileyi). Eddy. 1957. p. 162. fig. 406 
(coloration; Railroad Valley; "Railroad Valley spring- 
fish"). Moore. 1957. pp. 150-151, fig. 2-82 (color; 
teeth; Railroad Valley; "Railroad Valley springfish"). 
Frantz, 1958. p. 7 (in lake and steam survey, fide 
La Rivers, 1962, p. 517). Miller, 1958. p. 206, figs. 



228 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



15. 1(1 (relict habitat). Baii uv ct al. 1^)60. p. 21 
(freshwater; '■Railroad Valley killitish" ) . La Rivers. 
1962. pp. 27. 512. 517-520. Tigs. 231. 2.^2 ("Railroad 
Valley springfish": scientific-name and vernacular 
synonymies; original description quoted in lull; type 
locality; range; transplantation to near Sodaville. 
Nevada; taxonomy; compared with C. bailcyi: phys- 
ical data on habitat; associated fauna; newly hatched 
young in December). Uveno and Miller. 1962. p. 
522 (material studied). Bradley and Deacon. 1965. 
App. II, p. 2 (abundant. Railroad Valley). Hubbs 
and Hettler, 1965. pp. 245-24.S (physical habitat; 
tolerance). Hubbs. Baird. and Gerald. 1967. pp. 
104-115 (activity cycles related to diurnal fluctua- 
tions in dissolved oxygen; sound production; Duck- 
water Spring and Lockes Ranch Spring. Nevada). 
Cole, 1968, p. 477 (environmental factors, both 
species). Moore. 196S. p. 1(W (compared with C. 
bailcyi: Railroad Valley; "Railroad Valley spring- 
fish"). Bailey cI uI.. 1970. p. M) (freshwater; "Rail- 
road Valley killifish" ). 

As noted before (p. 71). the detaileci syste- 
matie ti-eatment of Crcnichthys and of C. ncvddcie 
is deferred. 

It seems virtually eertain llial Cicnichlhys 
iic\(iiUic has maintained existenee. as native, only 
in the warm .springs of Duekwater Valley, a flood- 
water arm of Railroad Valley, and in the warm 
springs on Loekes Raneh in the northwestern 
part of the main valley, both in Nye County, 
Nevada. Following are data on pre.served speei- 
mens of the species in the Museum of Compara- 
tive Zoology. Harvard University; University of 
Michigan Museum of Zoology; United States Na- 
tional Museum; University of Nevada. Las Vegas; 
Nevada State Museum; and Arizona State Uni- 
versity. 

MCZ 32'»4X (holotype). and LIMMZ '■)5()24 (paratype) : 
isolated warm spring at Duekwater. near north end ot 
Warm Spring Valley. 16 miles south and 46 miles 
west of Ely (originally misstated as 16 miles east and 
46 miles south of Ely — see La Rivers. 1962. p. 51t^). 
in 1. 12 N.. R. 56 E.; caught in a glass jar by Dr. 
and Mrs. C. T. Brues on July 21. \'>M). from the then 
abiuidant population; the holot\pe anil the parat\pe 
are maturmg females respectively 44 aiul 29 mm. in 
standard length. 

UMMZ 124941 and USNM 11750'); Wiirm Sprmg 
pool at source of Duekwater Creek, 5.2 miles by road 



above Duekwater. in T. 1.^ N., R. 56 E.; seined by 
Carl L. Hubbs and family on August 18, 1938, from 
the then teeming popiilation; 778 young to adult, of 
both sexes. 10-55 mm. long. 
UMMZ 1.^2176: spring and outlet, tributary to Duek- 
water Creek, beside Indian Colony, in T. 12 N.. R. 
56 E.; seined by Carl L. Hubbs and family on Sep- 
tember 9. 1934. from a teeming population; 1,677 
young to adult. 13-71 mm. long. 

UMMZ 132178; Duekwater Creek about 3 miles above 
Duekwater Store, in T. 12 N.. about on R. 55-56 
line; seined by Carl L. Hubbs and family on Septem- 
ber 8. 1934; 23 young to adult, 18-39 mm. long. 
UNLV 119: spring near highway at Duekwater; J. E. 
Deacon. Karl Larsen, and Ken Giles; August 31, 
1961; 166, 6-42 mm. long. 

UNLV 520 and ASU 3904: Duekwater; J. E. Deacon; 
June 5. 1964; 106. 19-54 mm. long. 

UNLV 552 and ASU 4124: Duckwaler; J. E. Deacon 
and M. B. Rheuben; .September 15. 1964; 64. 13-34 
mm. long. 

UNLV >-)46 and ASU 4218: Duekwater Spring; J. E. 
Deacon; October 2. 1966; 18. 27-51 mm. long. 

UMMZ 132173 and Nevada State Museum: large hot 
spring on Lockes Raneh in Railroad Valley, about a 
half mile from the ranch house, in T. 8 N., R. 56 E.; 
taken by derris by Carl L. Hubbv and family on Sep- 
tember 8. l')34. as a small portion of the teeming 
population; 1.158 young to adult of both sexes, 10- 
55 mm. long. 

UMMZ 132175: two other pools on Lockes Ranch; 
taken by Carl L. Hubbs and family on same day. by 
hand and small seine; 65 \oung to adult. 18-4! mm. 
long. 

UMMZ 181745: main hot spring on Lockes Raneh. ca. 
a hall mile northeast o\ main ranch house; seined by 
Robert Rush Miller and family on July 16. 1963; 319 
young to adult 15-4*^) mm. long. 

UNLV 261: ditch at Lockes Ranch; J. E. Deacon. 
Clark Hubbs. and Bernard J. Zahuranec; February 2. 
1963; 29. '■)-28 mm. long. 

UNLV 6()i) and .ASU 3905: Lockes Ranch; J. E. 
Deacon; June 5. 1964; 112, 14-51 mm. long. 

UNLV 905: Locke^ Ranch Spring; J. E. Deacon; 
October 2. 1966; 271. 11-40 mm. long. 

No evidence was found in our field work of 
1938 that Crciiiclitlnw lives in the various cool 
springs, on the east side of Railroad Valley, in 
which local forms of Cihi bicolor occur ( Hubbs 
and Miller, l'-M8b, p. 91 ). Nor have we found 
it, either then or in 1969, in Currant Creek, the 
eastern tributary to the valley, either in the lower 



VOL. Vll HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



229 



section, near the Currant Store, or higher in the 
creek. 

Fortunately for its perpetuation, since the 
species is so limited in habitat and distribution, 
Crenicbthys nevadae has been successfully trans- 
ferred outside the study area. The intentional 
transplant was thus recorded by La Rivers ( 1962. 
p. 518): 

.... Successfully transplanted by Tom 
[Thomas J.] Trelease into artificial ponds at 
Sodaviile, southeastern Mineral County on 
September 4, 1947. This natural seep of warm 
water on the west side of U. S. Highway 95 
was later enlarged by bulldozer action into a 
small pond. The transplant was motivated be- 
cause of the possibility that blackbass would 
be planted in the springfish's home locality. 
The f) original fish have become many. 

Having heard in 1 970 of the abundance of a small 
fish seemingly of this type about Sodaviile, Mil- 
ler and family checked the report on July 2, 1 970, 
and indeed found Crenichthys nevadae still abun- 
dant in the hot springs (36-38° C; air 34°), 
particularly in the lower of the two shallow pools 
that have been dug out. The pool at the head 
spring measured ca. 4 ;-^ 6 m. and the lower one 
ca. 6 X 6 m. Two more, quite small springs, with 
some springfish, open between the two pools, and 
the total outflow feeds into an elongate, deeper 
pond, with much vegetation, in which no fish 
could be seen. In the two pools containing the 
springfish the water was clear but very easily 
roiled: the bottom was of flocculent silt and hard- 
pan, with some gravel; the vegetation comprised 
green algae and. marginally. Sciipus and Jiinciis. 
The springs are across the highway from Sodaviile 
(American Mining Corporation). 

TRANSFERS OF GREAT BASIN FISHES 

(OTHER THAN GAME SPECIES) INTO. 

BETWEEN, AND FROM THE 

NORTH-CENTRAL BASINS 

Before considering the introductions of more 
exotic fishes into the north-central Great Basin, 
to enhance the sport fishery and for other pur- 



poses, we treat the highly probable to certain 
transferences by man of fish species endemic to 
the Great Basin into, between, and out of some 
of the 2 1 endorheic basins under study. We have 
found evidence for fifteen such transfers, involv- 
ing ten Great Basin fishes. The Utah chub, Gila 
atraria, has been introduced once, we suppose, 
into each of four among the 21 basins. Catosto- 
miis { Paiitosfeiis) platyrhynchus, Rliiiiichthys 
o.sciiliis robustus, Gila robiista jordani. Richard- 
soiiius egregins, Moapa coriacea. and Empetrich- 
iliys latos latos. all exotic to the 21 basins, have 
each been stocked once into a single basin. The 
same subspecies of Rhinichthys and the species 
Rclictiis solitarius have each been transported 
once from one of the north-central Great Basin 
endorheic units into another. Gila bicolor sub- 
species almost surely, Relicttis solitarius prob- 
ably, and Crenichthys nevadae certainly, have 
each been transferred into a basin outside the 
study area. Particulars are given or referenced 
below, under species headings. 

Because anomalous distributions have generally 
gone undocumented, and because they may be of 
considerable potential zoogeographical confusion, 
much effort has been expended, through exten- 
sive local inquiry, through examination of old 
maps, and otherwise, to arrive at a definite or 
highly probable conclusion as to whether each 
anomaly is attributable to a natural or an artificial 
event. Some of our conclusions rest on differential 
plausibility, and some reverse our former evalua- 
tion, for instance in regard to the native or intro- 
duced status of Catostomiis and Relictiis in Spring 
Valley. 

Catostomus (Pantosteus) platyrhynchus 

( Cope ) . 

Through a reappraisal of the evidence, we now 
attribute the occurrence of a mountain sucker 
{Pantosteus) in Spring Valley, the site of pluvial 
Lake Spring, to an introduction, rather than to the 
survival of a Glacial relict. In our early summary 
( Hubbs and Miller. 1948b, pp. 56-57) we men- 
tioned having heard that small fish occur in 



!30 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



"niinor springs forming a series on the east side 
of tlic valley,"" and we reported our collection of 
two species in Spring Creek, the northern axial 
tributary to the enclosed Spring basin. The re- 
port of fish on the east side of the valley referred, 
we now think almost certainly, to the introduced 
Utah chub, Gila atnu'ui. far to the south. One 
of the species collected in Spring Creek was 
Rcl'ulus solitariiis. for which we find evidence 
(pp. 233-235) that it was introduced into the 
basin. The other species was the mountain sucker. 

which we characterized as "a rare one a new 

sucker, with characters that line it up best with a 
Bonneville, or po.ssibly a Colorado species."" Still 
thinking the species to be distinct, one of us ( Mil- 
ler, 1^52, pp. 14, 28-29, fig. 13), after examin- 
ing a few similar specimens of doubtful prove- 
nance being sold in southern Nevada for bait, 
called it the '"dusky mountain-sucker, Puntoslciis 
species'" and stated that it is known only from the 
northern part of Spring Valley. It was so rare there 
in 1938 that we could collect by prolonged effort 
only three specimens, along with 1 ,073 of Rcliciiis: 
after the first sucker was taken, two hours of fur- 
ther .seining yielded only two more. This was at 
Collection 32 (p. 207) for Rclicliis, in a short, 
spring-fed section of Spring Creek, at the histori- 
cal old Stone House, in a strenuous effort to ob- 
tain more specimens just 21 years later, at the 
same place, we took 498 relict dace, but not a 
single sucker. Furthermore, Collections 30 and 
31 in springs issuing in the same stream bed, 
taken in 1964, comprised 80 and 99 specimens 
of Rclliiii.\ but not cine sucker. Nor has any other 
trace of suckers been found in Spring Valley (or 
in any of the other basins here treated ). 

The location of the Stone House on the old 
Overland Mail Route, later the Lincoln Highway, 
presumably facilitated a transfer of the sucker 
from some place in either the Bonneville or 
Lahontan watershed. The sucker as well as the 
dace may well have been brought in for bait or 
forage, for the flow was more ample when the 
senior author crossed this stream at the same 
place in the sunmier of 1915, on the retmn from 



participating in John O. Snyder's survey of the 
Bonneville fish fauna. 

Smith ( 1966, pp. 58-72. fig. 13) synonymized 
PaiUiistcus as a subgenus with Catostomiis, and 
synonymized the species previously recognized 
from the Lahontan system, 'lahontan,' with the 
one, 'plalyrhyncluis.' of the Bonneville .system 
( perhaps under-stressing considerable differences: 
he also synonymized 'ilclphiniis of the upper 
Colorado River system and 'jorclani' of the Great 
Plains with 'platyrhynchiis' ). Thus, it may not 
be feasible to determine if the mountain suckers 
were brought in from the west or from the east. 
Better agreement of the Spring Valley specimens 
in number of vertebrae with samples from the 
Humboldt River system than with tho.se from the 
Sevier River drainage ( Smith"s fig. 7, pp. 30-31 ) 
is a slight indication of introduction from the 
west. Our recognition of a distinct species for 
the Spring Valley fish, and the proposed vernacu- 
lar of "dusky mountain-sucker,"" were based 
largely on the intensity and wide spread of the 
dark color, which may well have been directly 
induced by the occurrence of the fish, at the time 
of capture, in extremely dense vegetation. 

Rhinichthvs osculus robustus ( Rutter). 

There is documentary evidence (pp. 107-109) 
that this species was twice introduced into Ruby 
Lake by the Nevada Fish and Game Commission 
to serve as forage for the largemouth bass, 
Microplcrii.s sahnoidcs. the object of a local sport- 
fishery. The first introduction, in 1950, was from 
a headwater of Humboldt River; the second, in 
1951. from "Sadler"s Ranch"" {i.e.. Big Shipley 
Spring) in Diamond Valley. There is indication 
of a very local and limited establishment, from 
the first stocking only. The first transfer was from 
outside the basin complex here under special 
treatment: the second, between two of the 21 
endorheic miits under study. 

Richardsonius egregius (Girard). 

"Red-striped shiners."" obviously of this species, 
were included with speckled dace in the 1950 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



231 



transfer, just recounted, from a headwater of 
Humboldt River into Ruby Lake. Apparently this 
stocking has not survived, nor has this Lahontan 
species managed to persist in any of the lake 
basins under treatment that formerly discharged 
into Humboldt River. 

Gila atraria ( Girard ) . 

The Utah chub, which stands in ill repute 
among fishery biologists because of its tendency 
toward population explosion and habitat domi- 
nance in artificial impoundments, has, we con- 
fidently believe, somehow gained entrance from 
its native waters of the Bonneville system into 
four of the 21 basins that are included in the 
present study. These intrusions have been into 
springs at Shoshone in Spring Valley, in the basin 
of pluvial Lake Spring (p. 64); into springs at 
Geyser Ranch in Lake (Duck) Valley, in the 
basin of Lake Carpenter (p. 66); into springs 
at Murphy Ranch in Steptoe Valley, in the basin 
of Lake Steptoe (p. 58); and into Comins Lake, 
in the basin of Upper Lake Steptoe (p. 60). 

The assumption that the stock of Utah chubs 
found persisting in Spring and Lake valleys re- 
sulted from introductions is complicated by the 
circumstance that these valleys are separated from 
upper Snake Valley by a low, flat sill, on the east 
side of which, not far distant, Utah chubs occur, 
presumably as natives, in Big Springs and the 
adjacent slough south of Garrison, Utah, in the 
pluvial drainage basin of Lake Bonneville. It 
might be theorized than an ancient outlet into the 
Bonneville system, or .some other stream connec- 
tion, could have led to the penetration of this 
chub into Spring Valley. However, there is no 
definitive evidence of such a discharge (p. 63), 
and no other indication of the occurrence of 
Bonneville fishes in Spring Valley. Even the 
trout was almost surely introduced (p. 64). Fur- 
thermore, the low passes would have facilitated 
the short-distance transport of chubs by man 
into Spring and Lake valleys, particularly during 
the years of early settlement. Indeed, as has al- 
ready been briefly noted, by Miller and Alcorn 



( 1 946, p. 1 82 ) , by us ( Hubbs and Miller, 1 948b, 
pp. 55, 57. 100), by La Rivers and Trelease 
(1952, p. 116), and by La Rivers (1962, p. 
397 ), direct testimony has indicated the introduc- 
tion of the Utah chub into the springs at Shoshone 
and at Geyser Ranch. On August 22, 1938, John 
Yelland told the senior author at Ely, Nevada, 
that this fish was almost certainly introduced by 
the original Mormon settlers into both sets of 
springs. This obviously well informed, well pre- 
served, and reliable man came into Steptoe Val- 
ley in 1881 and soon moved into Spring Valley. 
He volunteered that the chubs were stocked from 
Utah into the springs at Shoshone, where .settlers 
named Swallow came in from Fillmore, Utah, in 
the 1870"s or earlier. The old homstead in T. 1 1 
N., R. 67 E.. is still known as Swallow Ranch, 
and the Yelland Ranch is in T. 16 N., R. 68 E. 
Mr. Yelland added that the region about Geyser 
was settled about 1888 by pioneers, from Utah, 
named Edwards, Redheffer, and Wintergreen. 

It is even po.ssible that Indians may have 
brought Utah chubs into Shoshone Spring. In 
his "Nevada Place Names" Leigh ( 1964) stated, 
under "Shoshone," that the postoffice was given 
this Indian name when it was established in 1896 
for the Swallow Ranch, which was an ancient 
rancheria "in the head of Spring Valley, southerly 
from Wheeler Peak." 

As noted in the four articles cited above, 
respectively on pp. 182, 55, 116, and 387. we 
found another population of Gila atraria at Mur- 
phy Spring, in Steptoe Valley, where this species 
appeared to have been recently established, along 
with other fish, including the predatory Archo- 
pliies, which, together, seem to have replaced the 
native relict dace (p. 58). That occurrence, 
furthermore, seems too isolated to suggest in- 
digenity. It seems plausible that the chubs were 
brought there for forage. 

Unexpectedly, in August, 1969, Miller found 
Gila atraria swarming in Comins Lake. Steptoe 
Valley, under conditions even more conclusively 
indicative of introduction (p. 60). On inquiry. 
Thomas .f. Trelease, Chief of Fisheries for the 



9^0 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Nevada Department of Fish and Game, indicated 
inability to find any record of the stocking of 
tiiis fish, and thought that its cstabhshmcnt prob- 
ably resuhed from introduction of bait. Or, the 
chubs may have been stocked to supply forage 
for the well established gamefish. Local testimony 
indicated that a Mr. Swallow formerly operated 
the CCC (now C-B ) Ranch nearby, in the bend 
of Steptoe Creek, and it is to be recalled that an 
early settler, at Swallow Ranch in Spring Valley, 
next eastward, was named Swallow, and that the 
early Mormon inhabitants in that area are thought 
to have brought in Utah chubs, which we found 
there in abundance (pp. 64. 231). These cir- 
cumstances suggest that the introduction may 
have come from near Shoshone, the site of the old 
Swallow Ranch. 

Gila robusta jordaiii Tanner. 

The Pahranagat chub was stocked on Septem- 
ber 26, 1972. in one of the spring-fed refugium 
ponds recently constructed at Shoshone, on the 
east side of the southern part of Spring Valley. 
Nevada, as part of an effort to insure the survival 
of threatened species and subspecies of desert 
fishes (James Yoakum and Dale V. Lockard, per- 
sonal communications, November, 1972). This 
subspecies was described by Tanner (1950) as 
Cilii jonliini and has been discussed by La Rivers 
(1962, pp. 393-395, fig. 188) under the sub- 
species designation, in agreement with our ap- 
praisal. It is confined to certain springs and 
spring-fed creeks in Pahranagat Valley, in the 
course of pluvial White River. The transplant 
comprised 20 young specimens, which had been 
caught on the same day in the stream course be- 
tween the outlets of Crystal and Ash springs. 
Nearly a month later some were still living in the 
pond. 

Gila bicolor Girard. unnamed subspecies. 

The most widespread of the several subspecies 
of Gild hicolor inhabiting the basin of pluvial 
Lake Railroad (pp. 36. 142) has been found 
to occur also in Stone Cabin Valley (also known 



as Willow Creek Valley), just to the southwest, 
across a definite divide. The discovery of chubs 
here came as a distinct surprise, for this valley 
had been included by us in what we had treated 
(Hubbs and Miller, 1948b, pp. 45-51) as the 
Area of Sterile Basins, totally lacking in native 
fish. Having heard that a minnow was common 
there, the Hubbses. and Boyd W. Walker, collected 
tui chubs at Stone Cabin Ranch. 55 km. east- 
northeast of Tonopah, in 1946. They occurred 
there in the spring waters, in extreme abundance 
in one pothole spring with flocculent bottom, close 
to the ranch headquarters. The circumstances 
clearly suggested an introduction. Local testi- 
mony then indicated that the chubs, which had 
been present for several years, had been brought 
in from Twin Springs, in Hot Creek Valley [only 
40 km. due east, and there is a direct road con- 
nection]. Later comparison of the profuse ma- 
terial from each place showed that the specimens 
from Stone Cabin Ranch (H46-52: UMMZ 
144450) agree nicely with those from Twin 
Springs. Testimony of Forest Ranger Crane, in 
1938. that no native fish occur in Stone Cabin 
Valley (p. 19) is consistent with our findings. 

Moapa coriacea Hubbs and Miller. 

The Moapa dace was stocked in another one 
of the refugium ponds at Shoshone, on the same 
day and for the same purpose as Gila robusta 
jonlani was transferred ( same personal com- 
munications ). The genus and species were de- 
scribed by Hubbs and Miller ( 1948a, pp. 1-14, 
28. pi. I, fig. 1 ) as endemic in the Moapa River, 
a tributary of Colorado River, in the lower course 
of pluvial White River. The transplant of this 
species also comprised 20 individuals, which 
were half-grown to adult. Some were seen alive 
in the refugium nearly a month later. 

Relictus solitarius Hubbs and Miller. 

Two distributional circumstances suggest the 
possible transference of the relict dace between 
minor pluvial basins. We interpret the occurrence 
of the species in the adjacent Franklin-Gale and 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



233 



Waring-Steptoe pluvial lake systems as having 
resulted from an ancient stream transfer (p. 43). 
The finding of the species in Spring Valley, on 
the contrary, is not plausibly so explainable, as 
we have noted ( 1948b, p. 57). We are convinced 
that the occurrence of this species, as well as of 
Catostomus phifyrhynclius and Gila atraria. in 
this valley, is attributable to introduction by man. 
One of the reasons why we think the relict 
dace was introduced into Spring Valley was thus 
expressed by us (1948b, p. 56): "Most of the 
creeks and springs in the valley [Spring Valley] 
seem to be devoid of native fish life, and Cope 
and Yarrow ( 1875 ) marvelled at the lack of fish 
in this valley (then named Schell Creek Valley)." 
This was stated, obviously by Cope, in the brief 
account of "Siboma atraria var. longiceps. Cope" 

as follows (on p. 668): 

I 

This variety was discovered by Dr. H. C. 
Yarrow in Snake Creek Valley, Nevada, who 
remarked that, while very abund;int and the 
only species of this creek, in Schell Creek 
Valley, not far distant, no fishes whatever are 
found. The conditions of life being apparently 
similar in both streams, their difference in this 
respect was not explained. 

In view of the import of this statement on the 
original native distribution of the relict dace, a 
search was made of the early cxplorational reports 
and the early maps of the Great Basin, covering 
the third quarter of the last century, to try to fix 
the locale of Dr. Yarrow's "Schell Creek Valley." 
This search failed to verify our belief that his ob- 
servations can be attributed definitely to Spring 
Valley. In fact, the early reports and maps seem 
to have placed this valley and its creek variously 
along the old Overland Mail Route from the 
southwestern arm of Antelope Valley westward 
over the Spring Valley Creek section of Spring 
Valley to the northern part of Steptoe Valley ( as 
we use this term) — a distance of about 35 km. 

The least plausibly correct location of "Shell 
Valley" (listed by McVaugh and Fosberg, 1941. 
p. 17 ) is given in the narrative and accompanying 
map of Captain J. H. Simpson's report on his 



epochal exploration in 1858-59 to determine a 
wagon route from Camp Floyd in Utah to Genoa 
in Carson Valley (now in western Nevada). A 
critical reading of the narrative places the valley 
in the very arid southwestern arm of Antelope 
Valley, and the draftsman showed it imaginatively 
as a mountain-rimmed depression. The text de- 
scribes the basin "as a shallow valley, called 
Shell Valley, on account of it being covered with 
shale"! In Appendix A of the Simpson report the 
Mail Station near the present site of Schellburn 
is designated "Shell Creek, east side of Steptoe 
Valley, mail station" (this appendix may well 
have been added subsequent to the field work 
and initial narrative, for the publication of the 
report was delayed until 1876). 

Confusion is confounded by the map of Nevada, 
on a scale of 1 8 miles to the inch, that was in- 
cluded in "the Commissioner of the General Land 
Office, 1867, Report for the year 1866" (six 
years prior to Yarrow's trip, presumably on the 
same Overland Mail Route, and seven years after 
Simpson's exploration, but ten years before Simp- 
son's report was published ). An earlier large-scale 
map in the 1862 Commissioner's Report also 
names the valley, but not in a definitely identifi- 
able position. The I 866 map, which was rather 
poorly reproduced by Wheeler (1971, opp. p. 
108), shows the old Overland Mail Route, in- 
cluding "Capt. Simpson's Route" with the Ante- 
lope Spring, Spring Valley, Shell Valley, Shell 
Creek, and Egans Cation mail stations so well 
spaced as to make it appear that the "Shell Val- 
ley" station was in the valley of the present Spring 
Valley Creek, probably close to the old Stone 
House (the site of Collections 32 and 33 for 
Rclicliis solitdrius, here supposedly later intro- 
duced ), whereas the Shell Creek station was close 
to the present axial stream bed of Steptoe Valley, 
approximately at the present location of "Schell- 
burn," which name over the years has been vari- 
ously spelled by combining Schell- or Shell- with 
-bourne, -bourn, or -burn, and has been applied 
to a mail station, a ranch, a pass, and a fort, all in 
the same immediate vicinity. 



234 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Gibbes" map of California and Nevada, of 
1873, prepared, after the mining rusli was well 
underway, by a man who participated in the early 
surveys (p. 8), shows Schellburn, from which 
Schell Creek is shown flowing west to the 
"Slough," with '"Schell Valley" named from there 
northward to the "Sink" ( Goshute Lake ) : there- 
fore in the northern part of Steptoe Valley. Clark 
and Riddell ( 1920) also placed Schell Creek in 
the same position, and noted that it rises in a 
large spring on the pass. 

Obviously the name of Schell Creek has been 
applied over a rather wide area. In fact. "Schell 
Cr. Dfistrict]." is printed on the Rand, McNally 
map of 1881 (p. 8) above and well to the east 
as well as to the west of "Schellburn." which is 
located on a north-south road ( as on sc)me other 
maps ) . 

"Schell Creek Valley" may well have derived its 
name from the adjacent Schell Creek Mountains 
or Range, which presumably in turn derived the 
name from the minor stream known as Schell 
Creek ( perhaps standing for Schellbourne Creek, 
just as "Cleve Creek" stemmed Uom Cleveland 
Creek). Major geographical features in the gen- 
eral area were named for water sources, which 
in moister regions are relegated to insignificance. 
Sulphur Spring Range is another notable ex- 
ample, for the long and lofty range cwershadows 
the minor Sulphur Spring ( described under Loca- 
tion G6, p. 154). 

The vagaries of geographical nomenclature in 
the Great Basin, specifically in reference to Schell 
Creek and the Range, were thus expessed by 
Leigh ( 1964, pp. 54-55): 

Schell Creek Range is a long prominent 
range extending from the southern to the 
northern limits ol White Pine Coimty ... In 
the [lorth segment ol the range is Schellbourne 
Pass which was named after a small mining 
camp nearby. This camp, in the Cherry Creek 
district, was known at one time as Schell Creek 
[but Cherry Creek lies on the western side of 
Steptoe Valley, just north of Egan Canyon], 
and this cropped form, by extension, was ap- 
plied to the long important Range. Schell- 
bourne is, of course, a proper name. Schell 



Creek, near the Pass, was a relay station for 
the mail riders; soldiers were stationed there 
for protection of mail and travellers, and the 
place became known as Fort Schellbourne [on 
at least one modern map shown in the eastern 
part of the Steptoe Valley flat]. The cropping 
of the name from Schellbourne to Schell in 
early times was not sufficient to Americanize, 
i.e.. corrupt it; the name on modern maps has 
been redticed to Shell. This is an outstanding 
example of the gradual downgrading of a place 
name. 

There is, to conclude, no apparent way to de- 
termine the definite location of the stream in 
"Schell Creek Valley" in which Yarrow to his 
surprise found no fish. It may have been the 
present Spring Valley Creek, or it may have been 
Steptoe Creek (the "Slough"), in a portion of 
Steptoe Valley where the .stream is intermittent 
and may not have contained fish. There seem 
to be no other plausible alternatives. For evidence 
on the original ab.sence of the relict dace in Spring 
Valley we need to return to our own field work 
and to the experience of early settlers that was 
recounted to us in 1938. 

John Yelland, our convmcingly reliable in- 
former (p. 237), avowed that no minnows t^r 
other fish originally lived in Spring Valley. A 
specific reason for relying on his evidence is that 
he knew precisely where minnows occurred (and 
still occur) in the springs of Steptoe Valley. He 
said that he had seen every waterhole in Spring 
Valley from Shoshone Springs on Swallow Ranch 
(in T. 1 I N., R. 67 E. ), where he well knew the 
Utah chub, to Muncy Creek (in T. 20 N., R. 66 
E.). (His old Spring Valley homestead, still 
mapped as Yelland Ranch, is in T. 16 N.. R. 68 
E.) 

Bert Robison, a rancher native to the valley, 
testified to us on July 6, 1938. when he was 
middle-aged, that the deep pools and springs in 
the main part of Spring Valley, most of which 
dried up during the great drought of 1934, con- 
tained only carp iCypriniis corpio), "perch" 
(presumably Aichoplites inlcrruptus), rainbow 
trout {Salmo udirdncrii ), and "bass" (obviously 
Microptenis salmoidcs) — all introduced. He 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



235 



knew of "minnows" ( Relictiis soliUiriiis ) then oc- 
curring on his ranch ( at the old Stone House, 
where our Collections 32 and 33 were taken), 
and he, like John Yelland, knew of minnows in 
other springs in the north-central Great Basin, 
where we also knew of them. When on one oc- 
casion he drained one of the largest of the pools 
on Cleveland Ranch to eliminate carp prior to 
stocking trout, he found no minnows. He knew 
of no native fish in Shoshone, Worthington, or 
other springs along the base of Wheeler Peak. 
On the day of our interview we verified Mr. 
Robison's testimony by failing to see any fish dur- 
ing a careful examination of a number of the deep 
spring holes and tule-bordered pools in and below 
the hay meadows on the northern side of the large 
and well-watered Cleveland Ranch (near the west- 
ern margin of T. 16 N., R. 67 E. ) ; nor did we 
obtain any by seining two very deep pools. 

The sharply localized known occurrences of 
Relictiis solituriiis in Spring Valley (pp. 207- 
208) are inconsistent with its widespread and 
swarming abundance (except where locally sub- 
jected to major disturbance or predation) in the 
valleys where it seems surely to be native. Spring 
habitats that are profuse in Spring Valley, particu- 
larly but not exclusively along the western side 
of the northern part of the valley, at the base of 
towering Schell Creek Range, seem comparable 
to those abundantly occupied by the species in 
Butte and Steptoe valleys. Circumstances seem 
consistent with the hypotheses that the species was 
transferred from Steptoe or Butte Valley to Spring 
Creek at historic old Stone House, as noted in 
the accounts of Collections 30-33, and that the 
single population found (Collection 34) in a 
valley spring on the floor of Spring Valley 
stemmed from the established stock in Spring 
Creek through a tremendous flood ( p. 207 ) . Two 
bits of evidence favor the view that the stock of 
Relictiis came from Butte Valley: ( I ) the old 
Overland Mail Route (p. 233) ran from Spring 
Valley Creek across Steptoe Valley to Egan Can- 
yon along a course that has not yielded collections 
of Relictiis ( see fig. 1 4 ) and Steptoe Creek is. 



and was, impermanent, whereas the route from 
Egan Creek to Ruby Valley (see figs. 12 and 
14) ran by springs abounding in the relict dace; 
and ( 2 ) the low scale and gill-raker counts for 
the Spring Valley .series agree better with those 
for Butte Valley than with those from Steptoe 
and Waring valleys (p. 224; tables 43, 45). We 
thus conclude that Relictiis solituriiis has been 
introduced into Spring Valley, probably from 
Butte Valley. 

There is evidence that Relictiis solitariiis was 
once transferred to a spring in the Bonneville sys- 
tem of Utah (p. 226). 

Empetrichthys latos latos Miller. 

This, the only surviving subspecies of the 
Pahrump poolfish, a cyprinodont, was stocked in 
April, 1972, in one of the spring-fed refugium 
pools recently constructed at Shoshone, on the 
east side of the southern part of Spring Valley, 
Nevada, as part of an effort to insure the survival 
of threatened species and subspecies of desert 
fishes (James Yoakum and Dale V. Lockard, 
personal communications, November. 1972). 
This subspecies, along with the species, was de- 
scribed by Miller ( 1948, pp. 103-104, pi. 11), 
whose account has been quoted by La Rivers 
( 1962, p. 527. fig. 236). It still exists, very pre- 
cariously, in a single spring (Manse Spring) in 
Pahrump Valley. Nevada, within the pluvial 
drainage system of Lake Manly ( of Death Val- 
ley). The transplant comprised 15 individuals 
of varying size, from Manse Spring. Some were 
seen in the pond in October, 1972. 

Crenichthys nevadae Hubbs. 

The Railroad Valley springfish has been suc- 
cessfully slocked in warm springs in the Lahontan 
system (p. 229). 

INTRODUCTIONS OF EXOTIC FISHES 

A considerable number of exotic fish species, 
in addition to the mountain sucker and the Utah 
chub discussed above, have become established 



236 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



in the area of the north-central Great Basin under 
treatment. They appear to have contributed 
moderately to the food and recreational resources 
of the area, but have also played a role, varying 
from slight to catastrophic, in the depiction and 
even in the local extirpation of remnant native 
fish populations (see following section). In gen- 
eral, because they are members of a much more 
profuse biota, the introdticcd fishes are endowed 
with predatory and competitive capabilities far 
exceeding those of the few native fishes of the 
Great Basin, and of other western waters. A 
major compensation is the degree of adaptation 
the native fishes have attained, through prolonged 
and rigorous selection, to continued existence in 
habitats severely afflicted by desiccation and 
flooding. 

The north-central Great Basin has, happily, 
hardly been affected by the intriKluctions of tropi- 
cal home-aquarium fishes, such as have plagued 
some other parts of Nevada ( Deacon, Hubbs, 
and Zahuranec, 1964; Hubbs and Deacon, 1965; 
Minckley and Deacon, 1968), and other western 
slates. However, other exotic animals have been 
introduced into the area. One such is the bull- 
frog, Riiiiti catesbciuiui Shaw, which has been 
widely stocked, for food, in the Great Basin area, 
where it is regarded as a probable predator on 
native fishes (Miller and Hubbs, 1960, p. 28; 
La Rivers, 1962. pp. 434, 441, 475., 525). Most 
of the known establishments of bullfrogs in the 
Great Basin are southwest of the area under treat- 
ment, but one occurrence is near the northeastern 
corner of the study area, namely in Warm Springs 
of Independence Valley, the sole habitat of 
Rhinklilhys o.sculiiy letlioponis and Gila hicolor 
isohitd. Fortunately for the native fishes, two 
other widely spread exotics, naiuely the crayfish 
ProcdmhiiriLs cUirkii (Girard) and the mosquito- 
fish Gtunhusia affinis ( Baird and Ciirard). are 
known in the Great Basin largely or wholly from 
waters .south of the study area. 

The extent and status of the introductions of 
exotic fishes into Nevada have been investigated 
and discussed by Miller and Alcorn ( 1946). La 



Rivers and Trelease (1952), and La Rivers 
( 1 962 ) , and, for more southern parts of the state, 
by authors quoted above. To round out the ac- 
count of the fishes of the .study area, we add be- 
low, in systematic sequence, a few records and 
remarks, largely by cross references, to pertinent 
material listed in the text, as follows: 

h\ page references to listings of exotic fishies 
in the first part of this memoir, under the de- 
scriptions of the basins of the designated 
pluvial lakes ( pages listed below after the head- 
ing "Paleohydrography" ) ; and 

by serial Location number in the R series 
for the subspecies of Rhinichlhys osculus and 
m the Ci series for the subspecies of Gihi 
hicoltir: and by serial Collection numbers for 
Rcliclus .',^>lil(lrit(s. 

In general, the exotic fishes reported, observed, 
or collected were luentioned on the field sheets 
and in the habitat descriptions based thereon. The 
species is usually stated or obvious, except for 
"trout."" Some additional data and discussions 
are presented, tending to show that certain basins 
in the north-central Great Basin area under study 
have been devoid of certain native fish species, 
at least during the historic period. 

"Trout" 

Paleoh>drograph\': pp. 19 (Monitor Valley), 31 
(not native in Clover Valley tributaries). 45 (Ruby 
Valley tributaries). 60 (Comms Lake). Mi (Wilson 
Creek in Lake Valley). 68 ( lllipah Creek in Jakes 
Valley). 70 (Pine Creek in Pine Valley). 

Location: Rl ( Crass Valley ). Collection: 14 (Butte 
Valley). 

Also in diseussion oi springs in Murph\ Ranch in 
Steptoe Valley (p. 2()h). 

All evidence that wc have encountered, both 
positive and negative, points to the assumption 
that all trout that have been found in any of the 
streams or lakes within any of the endorheic 
basins herein selected for study are represented by 
introduced stocks. Any population that by re- 
mote chance may have persisted anywhere in the 
study area would almost certainly have repre- 
.scntcd one of the various subspecies of Sulmo 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



237 



clarkii, and would in high probability have be- 
come modified or eliminated by interaction with 
introduced trout. 

Salmo trutta Linnaeus. 

Brown trout have been widely spread in 
Nevada (Miller and Alcorn. 1946, pp. 174-175; 
La Rivers and Trelease, 1952, p. 1 14; La Rivers. 
1962, pp. 318-323. 643-645). La Rivers listed 
the species from within our study area in Skull 
Creek, Grass Valley, Toiyabe Range; in Pine 
Creek, Monitor Valley. Toquima Range; in 
Illipah Creek, Jakes Valley, White Pine Range: 
and in three streams in Steptoe Valley and five 
streams in Spring Valley. 

Salmo clarkii Richardson, subspecies. 

Paleohydrography : p. 64 (Spring Valley). 

We have already noted ( Hubbs and Miller, 
1948b, p. 57) that; "As a result of his survey 
in 1869, Wheeler ( 1875: 54) remarked on the 
lack of fish in the mountain streams of the Great 
Basin, except in the Humboldt Range and to the 
east of Snake Range." Any trout that existed in 
the "Humboldt Range," then largely what is now 
the Ruby Range, were presumably on the west 
slope, in the Humboldt River system. We have 
found corroborative evidence in the History of 
Nevada (edited by Myron Angel, 1881 ), where 
it is stated ( on p. 649 ) : 

There are only two streams in White Pine 
County that have fish in them. In 1876 trout 
were placed in Cleveland [now "Cleve"] Creek, 
in Spring Valley, and have multiplied rapidly 
since, l^ehman Creek, which flows into Snake 
Valley and then sinks, also contains trout, and 
it is supposed [a probably erroneous supposi- 
tion] that the Mormons, who formerly oc- 
cupied a portion of the valley |a correct state- 
ment], placed them there. 

John Yelland indicated to the senior author on 
August 22, 1938 (see also p. 245 ), that the trout 
stocked in Cleve Creek prior to 1881 were pre- 
sumed to have come directly or indirectly from 
Trout Creek, in the Bonneville drainage basin in 



Juab County, Utah: and further claimed, as a 
result of much observation in early years, that 
no fish were native in Spring Valley and that the 
rapidly established population in Cleve Creek 
served to stock many of the mountain streams of 
Spring Valley. On this basis. Miller and Alcorn 
(1946, pp. 175-178) referred the Spring Valley 
introduced trout to Salmo clarkii utah Suckley, 
but La Rivers and Trelease ( 1952, p. 114) and 
La Rivers ( 1962, pp. 283-284, 298) have stated 
that the Utah subspecies presumably no longer 
persists in this valley in pure form, because sub- 
species identified as 5. c. lewisi (Girard) and S. c. 
lu'iishawi Gill and Jordan have also been stocked 
in east-central Nevada. However, a fine collec- 
tion of cutthroat trout ( 1 3 juveniles to adults, 
UMMZ 165771), taken by Ted C. Frantz on 
November 5, 1953, in Pine Creek on the east 
slope of Spring Valley, on the west face of Mount 
Wheeler, represents a population distinctively 
characterized by strong and numerous basi- 
branchial teeth (20-34 or more), large dorsal 
and anal fins, a long head, and a chunky caudal 
peduncle — quite different from cutthroat trout in 
eastern and southern Utah, Salmo clarkii utah, 
and from the Lahontan cutthroat trout. Presum- 
ably, the trout in Pine Creek are descendants of 
the cutthroat that were introduced into Spring 
Valley in 1876. reportedly from Trout Creek, 
Utah. Two specimens from Trout Creek in the 
University of Utah examined by one of us 
( R. R. M.) are indeed of the same type as those 
from Pine Creek. Two cutthroat trout from Leh- 
man Creek (UMMZ 141701, taken at about 
8,000 feet in 1938), on the east (Bonneville) 
slope of Mt. Wheeler, closely resemble the Pine 
Creek trout and represent the native, probably 
undescribed, subspecies. In 1960, 1969, and 
1970 Goshute Creek, tributary to Steptoe Valley, 
in the Cherry Creek Range, was stocked with the 
Pine Creek trout, after a cloudburst on August 5. 
1955 had scoured the stream, eliminating the 
progeny of the 25,000 Yellowstone cutthroat 
trout that had been planted in the creek on Sep- 
tember 1 1, 1953 (James Yoakum, personal com- 
munication. November, 1972). 



238 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Similar testimony published by Miller and 
Alcorn (1946. p. 175) specifies the transfer in 
1873 of trout ("5. c. hcnslunvi") across the 
Toiyabe Range from Reese River tributaries to 
Kingston Creek in Big Smoky Valley ( the site of 
pluvial Lake Toiyabe), from which that type of 
cutthroat trout was stocked in Grass Valley 
(pluvial Lake Gilbert basin) and in Stoneberger 
and Roberts creeks in the drainage basin of 
ancient Lake Diamond. 

More recently the Yellowstone cutthroat trout. 
5. f. Icwisi. as well as the Lahontan cutthroat 
trout. S. c. henshawi. has been extensively 
stocked in mountain streams across Nevada ( Mil- 
ler and Alcorn. 1946. pp. 175-178; La Rivers 
and Trelease. 1952. p. 114; La Rivers. 1962. 
pp. 14. 18. 25. 31. 64. 90. 123. 138. 174. 184. 
185, 275-300. 645-646). 

We have failed, in repeated efforts, to find any 
indication that trout were native in any of the 
mountain streams on the east slope of the lofty 
Ruby and East Humboldt ranges, or elsewhere 
in the drainage basins of pluvial lakes Franklin 
and Clover, though they were no doubt native in 
the longer, lower-gradient streams, and lakes, on 
the western slope, in the Humboldt River system. 
Robert J. Behnke (personal communication, 
1969) has accumulated confirmatory indications. 
The short, steep-gradient streams on the east side, 
in U-shaped valleys that held major montane 
glaciers almost unique in the Great Basin (Sharp, 
1938), were apparently unsuited to the main- 
tenance or survival of trout. This circumstance 
is clo.sely analogous to the lack of native trout 
in Owens Valley. California, in the formerly 
glaciated valleys on the eastern escarpment of the 
Sierra Nevada. 

We have similarly failed to locate from any 
source any evidence that trout are or have been 
native in the basins of any of the other pluvial 
lakes under study. 

Salmo aguabonila Jordan. 

Golden trout have been stocked in Nevada 
(Miller and Alcorn. 1946, p. 189; La Rivers and 
Trelease, 1952, p. 121 ; La Rivers, 1962, pp. 199. 



317-318). with reported '"good results'" in Ruby 
Mountain lakes in Elko County. Those lakes 
are in the Humboldt River drainage basin, but La 
Rivers indicated (p. 3 1 S ) that "officials hope to 
utilize this site as a stocking source for other 
waters in the mountains of Elko County." 

Salmo gairdnerii Richardson. 

Paleohydrography : pp. 18 (Bailey Ranch in Dia- 
mond Valley). ."^^ (near C'lirric in Cioshiite Valley), 
.sX (Sleptoe Valley). 

l-ocations: R9 (Grass Valley), GIO (Fish Creek 
.Springs). Collection: 19 ( near Carrie ). 

Also in discussion of springs on Murphy Ranch in 
Steptoe Valley (p. .sS) and of pools and springs in Spring 
Valley (p. 234). 

All three treatises on introductions into Nevada 
document the widespread stockings of rainbow 
trout in the state. 

Salvelinus fontinalis (Mitchill). 

Paloohydrographv ; pp. .ss (near Currie). 58 (Step- 
toe Valley). 

l-Ocations and Colleclions same as lor Sahuo gaird- 
ncrii. 

The extensive introductions into Nevada of 
brook trout are likewise specified in the three 
papers just mentioned. 



Oihi;r Spi£Cies 

Coiegonus clupeafoimis clupea£oimis 

(Mitchill). 

In 1880, during the orgy of rampant introduc- 
tions, 25,000 eggs of the Great Lakes whitefish 
were sent to Eureka, but no records of plantings 
seem available (Miller and Alcorn. 1946. p. 188). 

Thymallus signifer tricolor Cope. 

Cirayling were unsuccessfully stocked in Ruby 
Valley in the I94()"s (Miller and Alcorn, 1946. 
pp. 187-188; La Rivers. 1962, p. 200). 



Cyprinus carpio Linnaeus. 



Paleohydrography: pp. 4(i (Butte Valley). ."^S (Step- 
toe Valley), dh ((ieyser Ranch, l.ake Valley). 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



239 



Locations: R12 and Gil (Independence Valley), 
G9 (Warm Springs, Newark Valley). Collections: 14 
(Stratton Ranch in Butte Valley), 22 (Campbell Ranch, 
Steptoe Valley), 23 (Steptoe Ranch, Steptoe Valley). 

Also in discussion of pools and springs in Spring 
Valley (p. 234). 

The introduction of carp into Nevada, with 
early enthu.siasm and later disillusionment, has 
been chronicled by Miller and Alcorn ( 1946, pp. 
180-181), La Rivers and Trelea.se (1952, p. 
116), and La Rivers (1962, pp. 18-21, 448- 
453, 525). 

Carassius auratus (Linnaeus). 

Paleohydrography: p. IS (Big Shipley Spring in 
Diamond Valley). 

Location: R8 ( Big Shipley Spring). Collections: 22 
(Campbell Ranch), 23 (Steptoe Ranch). 25 (Dairy 
Ranch) — all three in Steptoe Valley. 

Also in discussions of Cherry and Schell creeks, Step- 
toe Valley (pp. 206-207). 

Establishments of goldfish have been recorded 
by Miller and Alcorn ( 1946, pp. 181-182), La 
Rivers and Trelease (1952, p. 116), and La 
Rivers ( 1962, pp. 454-456), from several places 
in Nevada, some of which we have confirmed. 

Ictalurus (Ameiurus) species. 

Location: G9 (Warm Springs. Newark Valley). 

This report on "catfish," presumably of this 
subgenus, and one of "bullheads" in "Lake Don- 
pah-gate" in Diamond Valley (p. 17), are the 
only ones we have obtained for the study area. 
We have collected bullheads elsewhere in Nevada, 
for example /. n. ncbiilosiis ( Lesueur ) from Hum- 
boldt River near Carlin (G3) and /. m. melas 
(Rafinesque) from that station and from Carson 
River near Fallon (GI). The establi.shment of 
these and other ictalurids in Nevada has been 
treated by Miller and Alcorn ( 1946, pp. 182- 
184), La Rivers and Trelease ( 1952. p. 118), 
and La Rivers (1962, pp. 18, 479-497, 647- 
649). For our study area. La Rivers (pp. 648- 
649 ) listed /. nebiilosiis from Flynn Pond in Dia- 
mond Valley, Bassett Lake in Steptoe Valley, and 
Warm Springs Pond in Newark Valley. 



Perca flavescens (Mitchill). 

The only report of the yellow perch being intro- 
duced into the present study area is one for 
Bassett Lake in Steptoe Valley (La Rivers, 1962, 
p. 649). 

Micropterus salmoides salmoides (Lacepede). 

Paleohydrography: pp. 45 (Ruby Valley), fiO 
(Comins Lake). 

Locations: R4 (Ruby Valley), R12 and Gil (In- 
dependence Valley). Collection: 20 (Cardano Ranch, 
Steptoe Valley). 

Also in discussion of Ruby Lake (p. 198), of springs 
on Murphy Ranch in Steptoe Vallex' (p. 2()fi). and of 
pools and springs in Spring Valley (p. 234), 

The introduction into Nevada of largemouth 
bass, all we assume of the slower-growing north- 
ern subspecies ( Bailey and Hubbs, 1 949 ) , has 
been detailed by Miller and Alcorn ( 1946, pp. 
185-186), La Rivers and Trelease (1952, p. 
119), and La Rivers ( 1962. pp. 24, 432, 554- 
560, 649-650). 

Lepomis macrochirus macrochirus Rafinesque. 

Bluegills, presumably all of the nominotypic 
northern subspecies, have been introduced into 
various parts of Nevada, particularly in the Hum- 
boldt River system and in Lake Mead, but in our 
area of special study we did not encounter any 
sunfish, and the only record we have found for 
that area is for Bassett Lake in Steptoe Valley 
(La Rivers. 1962. p. 651 ). 

Archoplites interruptus (Girard). 

Paleohydrography: p. 58 (Steptoe Valley). 

Collection: 24 (Grass Springs on Lusetti Ranch in 
Steptoe Valley ) . 

Also mentioned in discussion of pools and springs in 
Spring Valley (p. 234). 

The Sacramento perch, the only native west- 
ern member of the sunfish family. Centrarchidae, 
has become scarce in the Central Valley of Cali- 
fornia but has become common in some localities 
in the Great Basin, into which it was transplanted 
long ago (Miller and Alcorn. 1946. pp. 186-187; 
La Rivers and Trelease. 1952. p. 120: La Rivers. 
1962. pp. 17. 18, 21. 545-553, 651 ). 



240 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



For the area under special study. Miller and 
Alcorn (p. 187) reported that in August. 1938, 
the species was common "in a lowland slough at 
Lusetti Ranch, about 16 miles north of Ely." 
This report was based on local testimony received 
by the senior author at Ely on August 22. 1938, 
from the local game warden. Earl Mangum. who 
slated that he had transported many of these fish 
to other localities in Nevada, including No. I Well 
in Railroad Valley and Preston and Lund in 
White River Valley. In 1962, James E. Deacon 
(personal communication. 1965) found the 
species persisting in a larger spring, on Lusetti 
Ranch, where it apparently had extirpated 
Rc'lictus soliuiriiis, which held out in three smaller 
springs not containing the predaceous centrarchid. 

On August 22, 1938, John Yelland (see p. 23 1 ) 
indicated that this fish had succeeded marvel- 
lously well, even in the brackish lakes of Spring 
Valley. La Rivers ( 1962, p. 549) had informa- 
tion that the species had died in this valley during 
a severe drought. He added that there was a sub- 
stantial but stunted population in Ba.ssett Lake 
in Steptoe Valley. His list of localities for the 
species in Nevada ( p. 651 ) included Bassett Lake 
in Steptoe Valley and Little Meadow Lake in 
Spring Valley. 

Although this species has proved predatory on 
a native minnow in Nevada, it is itself vulnerable 
to predation, because it does not guard its eggs. 
It has proved to be resistant to high-salinity 
waters, in general as well as in Spring Valley, and 
it is being extensively and successfully stocked in 
such waters (McCarraher and Gregory, 1970). 

Pomoxis nigromaculatus (Lesueur). 

The only record of the black crappie found 
from our study area is that for Fish Creek in 
Little Smoky Valley (La Rivers, 1962, p. 652). 

SURVIVAL AND CONSERVATION 

The studies that we and an increasing num- 
ber of colleagues have been conducting on the 
hydrography and ichthyology of the Great Basin 
and other arid parts of the American West have 



highlighted the peril that the fishes of the remnant 
waters have faced and continue to face. Several 
species and subspecies that had maintained 
existence over the millennia since late Pleistocene 
time of more ample water have become extinct 
during the past few years through the deteriora- 
tion of their environment, and others are on the 
verge of oblivion. It was our dubious privilege 
of describing posthumously (Miller and Hubbs, 
1960) a species and a subspecies of peculiar min- 
nows of the Colorado River system that had 
failed to survive the perils they had met during 
the two decades between their discovery and 
their description. More recently, we have added 
another posthumous description, that of Rhiiiich- 
rliy.s (>sculii.\ rcliqiiiis (Hubbs and Miller, 1972, 
pp. 104-105), here treated. Several other endemic 
fishes of the arid West, and even one entire genus 
( Empvtrichthys ) . have become endangered, and 
are being recognized as such in the international 
and U.S. federal 'Red Books" (referred to by 
Miller, 1972). Some of the documentation has 
been provided by Miller and Hubbs, I960; Mil- 
ler, 1961, 1968, 1969, 1972: Deacon, Hubbs, 
and Zahuranec, 1964; Hubbs and Deacon. 1965; 
Minckley and Deacon. 1968; Bunnell. 1970; 
Deacon and Bunnell. 1970; Miller and Pister, 
1971; Miller, 1972, 1973; Pister. in press. 

It is true that mass extinction has been the 
rule over the nullcnnia of aridity, as is stressed 
above (pp. 74-76). It is, therefore, small 
wonder that some subspecies and species have 
succumbed to the stress inflicted by man on their 
fragile habitats, while others have reached the 
endangered category. However, it is also true 
that the survivors have become almost un- 
believably resistant to desiccation, and through 
speciation during isolation have to some degree 
actually increased in number of subspecies. The 
very persistence of certain species and their 
speciation have bestowed upon the remnant forms 
a particular value and interest. 

The peril of existence has accelerated as the 
miniscule habitats of the fishes of the north-central 
Great Basin have been degraded or at times have 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



241 



even been eliminated by human activities, notably 
by the overuse of the waters for agriculture — 
usually marginal or often submarginal. As we 
have repeatedly documented in this report, habi- 
tat deterioration and threat to survival have been 
effected by diversion of water, by ditching, even 
by piping the entire waterflow, by removal of 
vegetation, and, disastrously at times, by the 
lowering of the fossil ground water through over- 
pumping ('water mining'). 

Another ominous threat to the survival of the 
native fishes in the area under special treatment, 
and in other western regions, has been the wide- 
spread introduction of exotic fishes and other 
animals, including the frequently introduced and 
established bullfrog. Rami cateshckma. a prob- 
able fish predator. Most of these introductions 
have been of species native to the faunally rich 
ecosystems of the eastern United States, wherein 
competition and predation have been highly 
evolved, whereas the native fishes of the isolated 
and faunally depauperate western waters have 
largely lost their ability to compete and to evade 
predation. In addition it may well be that the 
isolated native fishes have not developed, or have 
lost, immunity to parasites and diseases that the 
introduced species disseminate; for example, a 
case of parasitism affecting a native fish of south- 
ern Nevada, CreniclUhys bciilcyi (pp. 227-228), 
has been attributed, with appropriate timing, to 
the introduction of exotic aquarium cyprinodonts 
(Wilson, Deacon, and Bradley, I960. 

The occurrences of exotic fishes are mentioned 
under the pertinent Locations for Rluiuchthys 
and Gila and under the Collections for Relictiis, 
and the introductions of the exotics are further 
treated in systematic order (pp. 235-240). Three 
species not native to any of the 21 pluvial lake 
basins under detailed study, namely Catostomus 
platyrhyiicbiis. Richardsoniiis egregiiis, and Gila 
atraria, are treated among interbasin transfers 
(pp. 229-235). Of these the first two have 
seemingly not persisted where stocked. Gila 
atidiia, the Utah chub, has become established 
in four of the basins under study (pp. 58, 60, 



64, 66), wherein it has, as in some other places, 
increased inordinately in abundance. 

Fortunately, the area of special study has 
largely escaped the establishment, purposely or 
otherwise, of tropical home-aquarium fishes, such 
as has taken place, at an ominously increasing 
rate, in the southern parts of Nevada and Cali- 
fornia. Our special area has apparently also 
escaped, at least largely, the establishment of the 
mosquitofish, Gamhiisia affinis (Baird and 
Girard), which Miller and Hubbs ( 1960, pp. 
22-28), Myers (1965), and others, have re- 
garded as constituting a significant danger to the 
native fishes, largely through the consumption of 
fry. 

Survival Status Of The Endemic Fishes 

We now briefly review, in systematic sequence, 
the survival status of each of the species and sub- 
species of endemic fishes of the north-central 
area of the Great Basin under present study. As 
noted in the preceding section on Transfers of 
Great Basin Fishes certain species of desert fishes 
exotic to the area under treatment have been 
stocked in refugiuni ponds in Spring Valley. 

Speckled Dace, Rhinichthys osculus. 

Although one of the three subspecies of 
speckled dace that we have described appears to 
have become extinct in the past few years, and 
the other two have been hovering on the brink, 
the species as a whole, as we interpret it, can not 
be regarded as endangered, for it lives in a multi- 
tude of habitats over a great part of the American 
West. Many other subspecies, some named and 
others not, are no doubt also threatened. 

Lahontan Speckled Dace, Rhinichthys 
OSCULUS ROBUSTUS. Some of the more or less 
distinctive local populations of R. o. lobustiis are 
probably endangered, but the subspecies, as we 
treat it, is of such common occurrence over such 
a broad area as not to be feared in danger. Of 
the local populations treated in this report, those 
of a single tiny spring in Carico Lake Valley and 
of Indian Creek in Crescent Valley (p. 106) 



242 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



(both outside the basins of detailed study), may 
well be in danger. Presumably, various popula- 
tions will survive in the Humboldt River head- 
waters. Of the five local populations in the basin 
of pluvial Lake Diamond, that of the warm spring 
on Potts Ranch was on the verge of extirpation 
during our examination in 1938 (as is detailed 
on p. 114) and may well be gone now. A very 
similar stock, however, has just been found per- 
sisting in the small stream fed by springs about 
Dianas Punch Bowl, in the same valley, a few 
miles to the southward, and it probably has been 
increasing in population because of ditching. 
We suppose that the Coils Creek, Big Shipley 
Spring, and Birch Ranch populations (pp. 114- 
115) still persist, though the stock in Big Shipley 
Spring in 1938 was apparently considerably re- 
duced by modifications of the environment and 
perhaps by the establishment of goldfish. A 
sanctuary at Birch Ranch might save not only 
the slightly modified local form referred to R. o. 
robustus but also the more notably differentiated 
local endemic of Gila bicolor ( pp. I 54- 156). 

Grass V.-iiLLEY Speckled DAci:./^///;v/r//7//)'5 
oscui.us RELIQUUS. The evidence seems sound 
that this very distinct subspecies, which we found 
in 1938 confined to the spring area in Grass Val- 
ley, has since become extinct (pp. 121-128, 
240). The factors seem to have been the agricul- 
tural use of the water and, particularly, the stock- 
ing of trout. The loss of this Ice Age relict is a 
tragedy. It was a relict of some Pleistocene time, 
probably prior to the last pluvial. 

if a residual population of the subspecies should 
be found, it would be highly desirable to provide 
a sanctuary for it, either in Grass Valley or else- 
where. 

Clover Valley Speckled Daci:, Rhinich- 
THYS oscuLUS OLIGOPORUS. Near extinctions of 
the two known populations of this rather well 
differentiated subspecies seem to have resulted 
from agricultural practices inimical to the dace 
and from the stocking of trout in the impounded 
waters. Some form of protection seems highly 
desirable. 



Independence Valley Speckled Dace, Rhi- 
NiciiTHYS OSCUI.US LETHOPORUS. This sub- 
species, more distinct than its Clover Valley 
cousin, has been shown (pp. 134-135) to be 
extremely rare and, fortunately, secretive in dense 
vegetation. It has probably been reduced to a 
low population by the impoundment of the spring 
water and the introduction of predators, including 
black bass, carp, and bullfrogs. Chances for the 
survival of this interesting dwarf dace would be 
greatly increa.sed if the predators could be re- 
moved from the spring area or if a sanctuary, 
preferably in the home area, could be provided, 
with a device to exclude reinvasion by bass and 
carp. The companion endemic, Gila bicolor 
isolalii. could be granted an asylum in the same 
enclosure, for they have long been living together. 
It is strongly urged that this project be undertaken. 

Tui Chub, G/z.-i iucolor. 

The multiple local forms that we refer to this 
species are so many and so widespread as to calm 
any fears that the whole species is in danger of 
extermination, even though various local forms 
are definitely threatened. The apparently in- 
creasing trend for the creek ( G. b. obcsa ) and 
lake (G. b. pectinijer) subspecies to hybridize, 
owing probably to a decrease of the available 
waters in both quantity and variety, renders de- 
sirable the perpetuation of some nearly pure 
types. 

Humboldt Creek Tui Chub, Gila bicolor 
OBESA. It hardly seems reasonable that this wide- 
spread, central type could be regarded as in 
danger, but some recent examinations of the 
Humboldt River (in 1969) indicated that it had 
dwindled markedly in abundance and in distribu- 
tion in comparison with the findings of either the 
Lahontan survey of 1915 (Snyder, 1917) or our 
main operations of 1938 and 1942. It is to be 
hoped that at least some populations, such as that 
of Bishop Creek, may be saved. The remarkable 
several-step cline of races in the relatively short 
Humboldt River is almost unique and would be 
valuable to preserve if at all possible. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



243 



The very distinct race of Sulphur Spring in 
Diamond Valley that we refrained from accord- 
ing subspecies rank seems strictly confined to the 
head spring pool (p. 1 54) and therefore hangs on 
a delicate thread — if indeed it still survives. The 
nearby springs should be checked to see if there 
are other surviving pockets, and at least one pop- 
ulation should receive protection, preferably in 
the home territory. Perhaps not quite so urgently, 
but still definitely, the spring head at Birch Ranch 
(pp. 115, 155) should be utilized as a competitor- 
free habitat for its endemic races of Rhinichthys 
oschIks robiistiis and Gila hicolor obesa. 

Newark Valley Tui Chub, Gila bicolor 
NEWARKENSis. This distinct subspecies presum- 
ably maintains some populations, in Newark Val- 
ley, though we have not rechecked them since our 
collections of 1934 (pp. 157, 160). A new sur- 
vey is definitely in order. The delicate balance 
that such isolated stocks face was emphasized by 
the report we received of the complete die-off, 
through the freezing of a marsh area a few years 
before, of a population that had existed between 
the two main Locations sampled. Some com- 
petitor-free sanctuary, preferably at the type Lo- 
cation (G7), seems desirable. 

Fish Springs Tui Chub, Giia bicolor 
EUCHILA. This noteworthy subspecies, which with 
G. b. nenarkensis represents postglacial specia- 
tion, is certainly also worthy of secure protection 
— if, as seems highly probable, it still exists, as it 
did in 1938. The region of the type and only 
Location for this relict definitely appears to be 
the proper locale for a sanctuary. 

Independence Valli;y Tui Chub, Gila bi- 
color ISOLATA. This additional distinct, residual- 
endemic subspecies, which was discovered in 
1965 and 1966 (pp. 134. 175-180) surely also 
deserves some form of adequate protection, along 
with its companion relict. Rliinichthys osculus 
lethoponis. 

Relict Dace, Relictus solitarius. 

Although examples of local decrease in abun- 
dance and even of occasional local extirpation 



were encountered in our field work of 1934-71, 
this remarkable ancient relict seems to be, at 
least for .some time, safe from complete extinction, 
in the .spring-fed waters that it inhabits in the 
pluvial drainage basins of lakes Franklin and 
Gale, in Ruby and Butte valleys, and of lakes 
Waring and Steptoe, in Goshute and Steptoe 
valleys. 

The one really catastrophic depletion of the 
populations of the relict dace has occurred in 
Ruby Valley proper, between our discovery of 
the species there in 1934. when it existed in vast 
numbers, and 1965-67, when it could be located 
in very small numbers in only a very few habitats 
that had not been penetrated by black ba.ss and 
other exotic, predatory gamefishes, which 
abounded in the ponds representing Ruby Lake 
(pp. 197-199). It seems very doubtful that the 
probably slightly differentiated form of Ruby 
Valley can be saved, though it would be grand if 
a supply could be located to stock a spring area 
kept free of fish-predator invasion by the in- 
stallation of a siphon outlet. 

Fortunately a goodly supply of Relictus prob- 
ably remains in the northern part of Butte Valley 
that has had pluvial and modern-flood connection 
with Ruby Valley. However, the local form there 
seems somewhat different ( though not regarded 
as subspecifically separable) and it would be pref- 
erable to retain separately representative stocks 
of both Ruby and Butte valleys (and even from 
the Lake Franklin and Lake Gale divisions of 
Butte Valley). It would also serve science and 
conservation if sanctuaries could be built and 
maintained for the Johnson Ranch, Phalan 
Ranch, and above-Currie stocks of the Lake 
Waring basin, and for at least one, preferably 
several, stocks of Relictus in the basin of pluvial 
Lake Steptoe. 

Testimonial evidence was received (pp. 202- 
207 ) of the former enormous, in part seasonal, 
abundance of Relictus in Steptoe Valley, and of 
occasional extirpations, as at Murphy Ranch, 
probably as the result of manipulations of the 
water supply and the introduction of trout and 



244 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Utah chubs. A few other instances of local 
elimination were indicated by testimony to have 
resulted from manipulation of water, from stock- 
ing with Sacramento perch. Archopliles (p. 239) 
and other predators, and from other causes. How- 
ever, the survival of this relatively uniform, 
unique species does not seem to be in imminent 
peril. 

Railroad Valley Springfish, Crenichthys 

NEVADAE. 

There have been some signs or at least fears 
of the depletion of this remarkable endemic relict 
cyprinodont, which is restricted to two areas in 
Railroad Valley (pp. 227-229). but we have no 
evidence of its being in. or approaching, the en- 
dangered category. Nevertheless, its reliance on 
very limited warm springs places it in a precarious 
status. Fortunately, another population has al- 
ready been successfully established by transplan- 
tation (p. 229). The species justifies a "rare" 
rating, because of its few, very small habitats. 
The probably precarious status of the isolated 
populations of Crenichthys hailcyi, the only other 
species of the genus, and the nearly complete 
extinction of the whole genus Empctriclitliys. the 
only close relative of Crenichthys, need be kept 
in mind in planning for the perpetuation of 
Crciiiclitliys iicvadae. 

Hopi-FUL Signs Of Protfction For En- 

DANGHRED SpECIFS Of DESERT FlSHES 

Fortunately, in the current wave of awareness 
of Ihe need for conservation, the plight of the en- 
dangered desert-spring fishes has become appreci- 
ated by federal and state fish and game depart- 
ments, by the federal bureaus of Land Manage- 
ment and Reclamation, by the National Park 
Service, and by other agencies, as well as by 
biologists and conservationists. The Desert Fishes 
Council, made up of governmental employees, 
conservation agencies, and the concerned public, 
has emerged as a potent force. Positive actions 
to avert exterminations have been initiated, and 



the future appears less ominous. Such action has 
included the establishment of sanctuaries, with 
control of trespass and construction of devices to 
prevent reinvasion of exotic fishes ( Miller and 
Pister, 1971 ); transplantations; and the prohibi- 
tion of the use of springs for aquarium-fish rear- 
ing. Other vital steps include action to prevent 
the mining of ground water nearby, the with- 
drawal of public lands and the purchase of springs 
from private owners; and the inclusion of threat- 
ened species on the 'Red Book" lists of endangered 
animals. 

ACKNOWLEDGMENTS 

In this four-decade study we have been favored 
with the aid of almost countless persons. Geologi- 
cal and biological colleagues who have shared 
their knowledge and experience with us in our 
Great Basin studies are in large part listed in our 
earlier, general report (Hubbs and Miller, 1948b, 
p. 120), in which we added a statement, still 
true: "'In numbers beyond the possibility of li.st- 
ing, local, state, and federal employees, ranchers, 
prospectors and townfolk gave freely, in the 
generous spirit of the West, of help, advice, and 
information." In the same report we listed aides 
who assisted in the field and laboratory activities, 
prior to 1948. 

We now select, for special mention, alphabeti- 
cally, certain persons who particularly assisted us 
in our studies in the north-central Great Basin. 

Robert .1. Behnke shared with us conclu- 
sions from his studies on the native distribution 
of trout in the north-central Great Basin; he 
donated the second collection of GHa hicolor 
isoliitii. and he sent the extensive class report on 
this form written by one of his former students, 
Stephen Harold Berwick. Robert E. Brown pro- 
vided specimens of and ecological data on a 
stock of Rliinichtliys osciiliis robiistns that he 
found in l'-)72 in a hot-spring outflow by Dianas 
Punch Bowl. In 1971. Donald R. Cain of 
the Ely District Office of the U. S. Bureau of 
Land Management provided two supplementary 
collections of Rcliclus solitarius, along with a 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



245 



very welcome suggestion for the establishment 
of a preserve for this unique fish. Ted M. 
Cavender has indicated that his unpublished 
osteological studies of "Hybopsis" crameri justify 
its continued segregation as a genus. Oregonich- 
thys (considered in the discussion of Relictus). 
Patrick Coffin, Nevada Department of Fish 
and Game, provided, with good data, the only 
known specimens of Rhinichthys osciilus robiistus 
from Carico Lake Valley. James E. Deacon, 
University of Nevada, Las Vegas, provided speci- 
mens that he collected in the area of study, along 
with ecological and other data; and he rendered 
other help. 

Lee R. Dice loaned a 2-foot Paulin al- 
timeter for our 1938 and 1942 trips. The second 
collection of Gila bicolor isolata was loaned 
by W. I. Follett (California Academy of 
Sciences). When he was a student, John 
Thomas Greenbank made counts and measure- 
ments for us on the many local forms of Gila 
bicolor. Thomas P. Lugaski, graduate student at 
the University of Nevada, Reno, shared some in- 
formation derived from his current studies of the 
fishes in north and central Nevada. Ira La 
Rivers. University of Nevada. Reno, has shared 
local testimony regarding the extermination of 
Rhinichthys osculiis reliqiiiis in Grass Valley, and 
has otherwise assisted us. When he was Refuge 
Manager at the Ruby Lake Wildlife Refuge, 
Donald E. Lewis obtained collections of minnows 
in Ruby Valley and located information about 
introductions. The Nevada Department of Con- 
servation and Natural Resources supplied copies 
of the water-resource reports of various valleys. 
Much information, advice, and encouragement 
has come from C. T. Snyder and associates of the 
United States Geological Survey in Menlo Park, 
California. Their proposal of the name "Lake 
Hubbs" is much appreciated. Robert H. Soulages, 
a biology student at University of California, 
Davis, who has extensively explored underground 
and surface waters in Nevada, has provided in- 
formation regarding physiography, waters, and 
fishes in Roberts and E"an creeks and elsewhere 



in the state. Thomas J. Trelease, Chief of Fish- 
eries, Nevada Department of Fish and Game, 
provided information on fish transfers and intro- 
ductions, granted collecting permits, and other- 
wise assisted us. For the expert preparation of 
the karyotype of Relictus solitariiis and for the 
pertinent interpretation and comparison, we are 
deeply grateful to Dr. Teruya Uyeno. 

Among the long-time residents of Nevada who 
recounted their memory of the introduction of 
fishes into some places and of their native occur- 
rence or ab.sence elsewhere — without which data 
the early record would be blank — we gratefully 
mention John Yelland. for his trust-inspiring in- 
formation on the introduction of trout (p. 237) 
and Utah chubs (p. 231). on the original habitat 
of the relict chub (p. 234). and on the presence 
or absence of other fishes. Another very informa- 
tive resident. Jerome Phalan Stratton (pp. 199, 
206. etc.), who grew up in Butte Valley, helped 
particularly on place names, on the habitats of 
the relict dace, and on the presence or absence of 
minnows in various basins. His information, like 
that of Mr. Yelland. is regarded as highly reliable, 
as it stemmed from long ranching experience, in- 
volving observations on the fish in various springs. 

Among others who helped with local informa- 
tion were ranchers Campbell, W. J. Gardner, 
Ellen Valiee. and Mrs. Zubiri, all in Steptoe Val- 
ley; William McGill of Ely, Earl Mangum ( local 
game warden at Ely), and Dale V. Lockard of 
the Nevada Fish and Game Department, all deal- 
ing with Steptoe Valley and the relict dace; Forest 
Ranger Crane, rancher George Potts, and gradu- 
ate student Robert E. Brown, in Monitor Valley; 
rancher Jorge Jacobsen in Diamond Valley; 
ranchers Isador Sara in Fish Springs Valley, Billy 
Moore in Newark Valley, Vernon Westwood in 
Clover Valley, Bert Robison in Spring Valley; 
and Mr. Moorman at Illipah Creek in Jakes Val- 
ley; also Seiar S. Hutchings of the Desert Range 
Station and Otto Fife of Beryl Junction. Utah, 
regarding Pine Valley. For other names see Index. 

James Yoakum of the United States Bureau of 
Land Manaeement, Dale V. Lockard. and other 



246 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



participants in tiie meetings of the Desert Fishes 
Council, have provided information on the efforts 
being made to preserve endangered fish species in 
tiie region under treatment. 

Frances Hubbs Miller typed and reviewed 
much of the initial manuscript, prepared the 
index, and helped in the field. Other members of 
our families have assisted, particularly in the field 
work. The final typing has been the task of 
Elizabeth Noble Shor, who has consistently pro- 
vided editorial suggestions. 

The maps and other figures accompanying this 
treatise were diligently prepared (from drafts by 
the senior author) by Martha B. Lackey of The 



University of Michigan and by Elizabeth Parker, 
David G. Crouch, and Howard G. Shirley of 
Scripps Institution of Oceanography. The photo- 
graphs of scales (fig. 46) and of fishes are the 
painstaking work of Louis P. Martonyi, of The 
University of Michigan. Precision planimeters 
were loaned by Professor Charles M. Davis, De- 
partment of Geography, The University of Mich- 
igan, and by James R. Moriarty. Scripps Institu- 
tion. 

The ichthyological researches of the authors 
have been generously supported by grants from 
the National Science Foundation, and from their 
respective institutions. 



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1971. Cave and spring fishes of the southern bend 
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1949. The black basses (.Microptcrus) of Florida, 
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Bailey, Reeve M., Ernest A. Lachner, C. C. Lini^sey, 
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1964. Nomenclature of the blue chub and the tui 

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BRADLE^, W. Glen, and James E. Deacon 

1965. The biotic communities of southern Nevada. 

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appendices. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



247 



Brues, Charles T. 

1932. Further studies on the fauna of North Amer- 
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Bunnell, Sterling 

1970. The desert pupfish. Cry California, the 
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1963. New data on the isostatic deformation of Lake 

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1970. Man and pupfish. Cry California, the Journal 
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Zahuranec 

1964. Some effects of introduced fishes on the 

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Deacon, James E., and Brian L. Wilson 

1966. Daily activity cycles of Crenichtbys bailcyi, a 

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1967. Daily activity cycles of Creiiichthys baileyi, a 

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Eakin, Thomas E. 

1960. Ground-water appraisal oi Newark Valley, 

White Pine County. Nevada. [Nevada De- 
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1961. Ground-water appraisal of Long Valley, White 

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1962. Ground-water appraisal of Diamond Valley, 

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Eakin, Thomas E.. Jerry L. Hughes, and Donald O. 
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1951a. Ground water in Rtiby Valley, Elko and 
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1951b. Ground water in Clover and Independence 
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Eddy, Sami'ei 

1957. How to know the Ireshwater fishes. Wm. C. 
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Engel. a. E. J. 

\9M. Time and the earth. Ameriean Scientist, vol. 
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Evans, Howard E.. and Eari E. Deubler. Jr. 

1055. Pharyngeal tooth replacement in Scinotilus 
utrdnuiculaUis and Clino.sUnnus cloiiinilus. 
two species of cyprinid fishes. Copeia, 
1955, no. I, pp. 31-11, figs. 1-5. 
Everett. D. E., and F. Eugene Rush 

1966. A brief appraisal of the water resources of 
Grass and Carico lake valleys. Lander and 
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1931. Physiography of Western United States. 
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534 pp.. 1 73 figs.. I map. 
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1061. A new map of Western Coterminous United 
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|0(i4. Review and annotated bibliography of ancient 
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Flint, Richard Foster 

1957. Glacial and Pleistocene geolog\. ,lohn Wiley 
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Frantz. Theodore C. 

105.S. "Railroad Valley ponds — Series A: 1-9, maps 
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GU KERT. ChARI ES H. 

1,S93. Report on the fishes of the Death Valley 
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CilRARD. Ch\RIES 

1.S56. Researches Lipon the cyprinid fishes inhabiting 



the fresh waters of the Mississippi Valley, 
from specimens in the museum of the 
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1968. Water-resources appraisal of Butte Valley, 
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Harrh I . J. R. 

1971. Water-resources appraisal of the Pilot Creek 
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HoPKiRK, John D., and Robert J. Behnke 

1966. Additions to the known native fish fauna of 

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1932. Studies of the lishes of the order Cyprino- 
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1940. .Speciation of fishes. American Naturalist, vol. 
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1041a. Fishes of the desert. The Biologist, vol. 22, 
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1041b. The relation of hydrological conditions to 
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1042. .Sexual dimorphism in the cyprinid fishes. 

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1967. La Jolla nalural radiocarbon LiicasLLrements V. 

Ratiiocarbon. vol. 0. pp. 261-294. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



249 



HuBBS, Carl L., and Walter R. Crowe 

1956. Preliminary analysis of the American cyprinid 
fishes, seven new, referred to the genus 
Hyhopsis, subgenus Erimystax. Occasional 
Papers of the Museum of Zoology — Uni- 
versity of Michigan, no. 578, 8 pp. 

HuBBS, Carl L., and Clark Hubbs 

1958. Notropis sciladonix, a new cyprinid fish en- 
demic in the Rio Salado of northeastern 
Mexico. Copeia, 1958. no. 4, pp. 297-.^07, 
figs. 1-3. 

Hubbs, Carl L., and Laura C. Hubbs 

1945. Bilateral asymmetry and bilateral variation in 
fishes. Papers of the Michigan Academy 
of Science, Arts, and Letters, vol. 30, for 
1944, pp. 229-310, figs. 1-2, pi. 1. 

Hubbs, Carl L., and Eugene R. Kuhne 

1937. A new fish of the genus Apocope from a 
Wyoming warm spring. Occasional Papers 
of the Museum of Zoology — University of 
Michigan, no. 343, 21 pp., 3 pis. 

Hubbs, Carl L., and Karl F. Lagler 

1964. Fishes of the Great Lakes region. The Uni- 

versity of Michigan Press, Ann Arbor, xv 
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1941. Studies of the fishes of the order Cyprino- 
dontes. XVII. Genera and species of the 
Colorado River system. Occasional Papers 
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1943. Mass hybridization between two genera of 
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1948a. Two new, relict genera of cyprinid fishes from 
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1948b. The zoological evidence/ Correlation between 
fish distribution and hydrographic history 
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States. //; The Great Basin, with emphasis 
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1965. Studies of cyprinodont fishes. XXII. Varia- 

tion in Lucania parva, its establishment in 
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a new species from an interior basin in 



Coahuila, Mexico. Miscellaneous Publica- 
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waters in the Great Basin of western North 
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Hubbs, Clark 

1970. Teleost hybridization studies. Proceedings of 
the California Academy of Sciences, Fourth 
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Hubbs, Clark. Ronald C. Baird, and Jerry W. Gerald 
1967. Effects of dissolved oxygen concentration and 
light intensity on activity cycles of fishes 
inhabiting warm springs. American Midland 
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Hubbs, Clark, and James E. Deacon 

1965. Additional introductions of tropical fishes into 
southern Nevada. The Southwestern Nat- 
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Hubbs, Clark, and George F. Drewr> 

1962. Artificial hybridization of Crcnichthys bailcyi 
with related cyprinodont fishes. Texas 
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110. 

Hubbs, Clark, and William E. Hettler 

1965. Observations on the toleration of high tem- 
peratures and low dissolved oxygen in 
natural waters by Crcnichthys hailcyi. The 
Southwestern Naturalist, vol. 9, no. 4, for 
1964, pp. 245-248. 

Illick. Helen J. 

1956. A comparative study of the cephalic lateral- 
line system of North American Cyprinidae. 
American Midland Naturalist, vol. 56. no. 
1, pp. 204-223. figs. 1-39. 

Jones, David T. 

1940. Lake Bonneville maps. Edwards Bros., Inc., 
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numbered]. 

Jordan, David Starr, and Barton Warren Evermann 
1896. The fishes of North and Middle America: a 
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America, north of the Isthmus of Panama. 



250 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Bulletin ot the United States National Mu- 
seum, no. 47. part 1. i\ + 1240 pp. 
Kafiikii, Takeichiro 

l^-'iS. Speciation in cyprinid fishes on the basis of 
intestinal differentiation, with some refer- 
ence to that among eatostomids. Bulletin 
of Freshwater Fisheries Research Labora- 
tory, Tokyo, vol. 8, no. 1, pp. 45-78, figs. 
1-20. pis. 1-8. 

KiMSEY. J. B. 

1954. The life history of the tui chub. SiplnilcUs 
hicolor (Girard). from Eagle Lake. Cali- 
fornia. California Fish and Game, vol. 40, 
no. 4, pp. .V)5-41(). figs. 1-8. 
King, Ci arence 

1878. Systematic geolog\ . United States Geological 
Exploration of the Fortieth Parallel. Wash- 
ington, vol. 1. \ii + 80.^ pp.. frontispiece. 
12 maps. 26 pis. 
King, Philip B. 

1958. Evolution ol modern siirlace features of 
Western North .America. //; Zoogeography, 
edited by Carl L. Hubbs. The American 
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Publication no. 51, Baltimore, pp. .^-60, 
figs. 1-11. 
Koi'EC, John A. 

1949. Ecology, breeding habits and young stages of 
Cicnichthy.s hailcyi, a cyprinodont fish of 
Nevada. Copeia. 1949, no. 1. pp. 56-61, 
figs. 1-2. 
La< iiNER, Ernest A., and Robert F. Jenkins 

1967. Systematics. distribution, antl c\cilution of the 
chub genus Nocaniis (C'vpnmdael m the 
southwestern Ohio River b.ism. with the 
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no. .\ pp. 557-580, figs. 1-7. 
Langiois, T. H. 

192'». Breeding habits of the northern dace. Ecology, 
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L\ Rivi Ks, Ira 

1952. A key to Nevada fishes. Bullctm of the 
.Southern California Academy ol Sciences, 
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1962. Fishes and fisheries of Nevada. Nevada State 
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782 pp., 270 figs., 1 map. 
La Ri\i rs. Ira, and T. J. Trelease 

1952. .'\n annotated check list ^^^ the fishes of 
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Li iGii, Reiflis Wood 

1964. Nevada place names / Their origin and sig- 
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Llislorical Society, Las Vegas / Lake Mead 
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LoNGWELL, Chester Ray, Richard Foster Flint, and 
John E. Sanders 

1969. Physical geology. John Wiley and Sons. Inc., 

New ^■ork, 685 pp., many figs.. 2.^ pis. 

Lugaski. Thomas 

1 972. ,\ new species of speckle dace from Big Smoky 
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vada Occasional Papers, no. .^0. 8 pp., 2 figs. 

MacArtirir, Rohert H., and Edward O. Wilson 
1967. The theory of island biogeography. Mono- 
graphs in Population Biology, Princeton 
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60 figs. 

Maxe^, G. B.. and 1. E. Fakin 

1951. Ciround water in Railroad, Hot Creek. Reveille, 
Kawich. and Penoyer valleys, Nye. Lincoln, 
and White Pine counties, Nevada. .State of 
Nevada Water Resources Bulletin No. 12 
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4. 5. 

McCarraher. I). B.. and Richard W. Gregor'i 

1970. Adaptability and current status of introduc- 

tion of Sacramento perch. Anhoplilcs iiiter- 
niptiis. in North America. Transactions of 
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4. pp. 700-707. fig. I. 
McVaugh. Rogers, and F. R. Fosberg 

1941. Index to the geographical names of Nevada. 

Contributions toward a flora of Nevada, 
Work Projects Administration of Nevada, 
no. 2'^). 216 pp.. I map. 
Merriam, C. W., and C. A. Anderson 

1942. Reconnaissance survey of the Roberts Moun- 

tains, Nevada. Bulletin ol the Geological 
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figs. 1-.^, pis. 1-4. 
MiLiER, Robert Rush 

1945a. Siiydcricluhys, a new generic name lor the 
leatherside chub o'i the Bonneville and upper 
Snake drainages of western LJnited States. 
Journal of the Washington Academy of 
Sciences, vol. .^5. no. 1. p. 28. 

1945b. A new cyprinid fish from southern Arizona, 
and Sonora, Mexico, with the description 
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104-1 II). pi. 1. 

1946. Ciirrclation between fish distribution and 
Pleistocene hydrograph\ in eastern Cali- 
fornia and southwestern Nevada, with a 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



251 



1948. 



195: 



1957. 



1958. 



1961. 



1968. 



1969. 



1972 



1973. 



Miller, 
1946. 



Miller, 
1960. 



map of the Pleistocene waters. Journal of 
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Bait fishes of the lower Colorado River from 
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Utilization of X-rays as a tool in systematic 
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Origin and affinities of the freshwater fish 
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187-222. figs. 1-19. 

Man and the changing fish fauna of the 
American Southwest. Papers of the Mich- 
igan Academy of Science, Arts, and Letters, 
vol. 46, for 1960, pp. 365-404, fig. 1. 

Records of some native freshwater fishes 
transplanted into various waters of Cali- 
fornia, Baja California, and Nevada. Cali- 
fornia Fish and Game, vol. 54, no. 3, pp. 
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Freshwater fishes. Vol. 4 Pisces, of Red Data 
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Threatened freshwater fishes of the United 
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Two new fishes, Gila hicolor snyderi and 
Catostoimis fiimeivenlris. from the Owens 
River basin, California. Occasional Papers 
of the Museum of Zoology — University of 
Michigan, no. 667, 19 pp.. 9 figs. 

Robert Rush, and J. R. Alcorn 

The introduced fishes of Nevada, with a his- 
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the American Fisheries Society, vol. 73. for 
1943 (■T945"), pp. 173-193. 

Robert Rush, and Carl L. Hubbs 

The spiny-rayed cyprinid fishes ( Piagopterini ) 
of the Colorado River system. Miscella- 
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Zoology — University of Michigan, no. 115, 
39 pp., 2 figs., 3 pis. 
1969. .Systematics of Gasterosteiis aculeatus. with 
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introgression along the Pacific Coast of 
North America; a commentary on a recent 
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69, figs. 1-2. 

Miller, Robert Rush, and Clark Hubbs 

1962. Gila pandora, a cyprinid new to the Texas 
fish fauna. The Texas Journal of Science, 
vol. 14, no. I, pp. 111-113. 

Miller, Robert Rush, and E. P. Pister 

1971. Management of the Owens pupfish, Cyprino- 
tlon radiosus. in Mono County, California. 
Transactions of the American Fisheries So- 
ciety, vol. 100, no. 3, pp. 502-509, figs. 
1-5. 

MiNCKLEi. W. 1 .. and James E. Deacon 

1968. Southwestern fishes and the enigma of "en- 
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3822, pp. 1424-1432. figs. 1-3. 

Moore, G. A. 

1957. Fishes. /// W. F. Blair ft <//., Vertebrates of 
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1968. Fishes. In W. F. Blair et al.. Vertebrates of 
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Morrison, Roger B. 

1965. Quaternary geology of the Great Basin. In 
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Morrison. Roger B.. and H. E. Wright 

1967. Quaternary soils. University of Nevada Desert 
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Myers, George S. 

1965. Ganihiisla. the fish destroyer. The Tropical 
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Nolan. T. B. 

1943. The Basin and Range province in Utah. 
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Pister, E. 
In press. 



Desert fishes and their habitats. 



252 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Reno, Harlev VV. 

1969. Cephalic lateral-line systems of the cyprinid 

genus Hyhopsis. Copeia, 1969. no. 4, pp. 

736-773. figs. 1-32. 
RiVAS, Luis Rene 

1964. A reinterpretation of the concepts of "sym- 

patric" and "allopatric" with proposal of 
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topic." Systematic Zoology, vol. 13, no. 1, 
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Rush, F. Eugene, and Thomas E. Eakin 

1963. Ground-water appraisal of Lake Valley in 
Lincoln and White Pine counties, Nevada. 
Ground-water Resources-Reconnaissance Se- 
ries, Report 24, 29 pp., 5 figs., 1 pi. 

Rush, F. Eugene, and Duane E. Everett 

1966. Water-resources appraisal of Little Fish Lake. 
Hot Creek, and Little Smoky valleys, Ne- 
vada. Nevada Department of Conservation 
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5 figs., 1 pi. 

Rush, F. Eugene, and S. A. T. Ka/mi 

1965. Water resources appraisal of Spring Valley, 

White Pine and Lincoln counties, Nevada. 
Nevada Department of Conservation and 
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Reconnaissance Series Report 3.^, 3(i pp., 
7 figs., 1 pi. 
Rush. F. E., B. R. Scott. A. S. Van Denburgh, and B. 
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1971. State of Nevada Water Resources and Inter- 
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Russell, Israel Cook 

1885. Geological history of lake Lahontan / A 
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Monograph of the United States Geological 
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RUTTER, Cl OlIDSLEV 

1903. Notes on fishes I rom streams and lakes of 
northeastern California not tributary to the 
Sacramento basin. Bulletin of the United 
States Fish Commission, vol. 22. for 1902. 
pp. 143-148, 2 figs. 

1908. The fishes of the Sacramento-San Joaquin 
basin, with a study ol their distribution and 
variation. Bulletin ol the United States 
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103-152. figs. 1-4, pi. 6. 



Schultz, Leonard P. 

1936. Keys to the fishes of Washington. University 
of Washington Publications in Zoology, vol. 
2, no. 4. pp. 105-228. figs. 1-50. 
Schultz, Leonard P.. and Carl L. Hubbs 

1961. Early nomenclatural history of the nominal 
cyprinid genus Oicgonichlhys and of the 
blennioid, P/iolis schulizi, fishes of western 
North America. Copeia. 1961. no. 4, pp. 
477-478. 
Schultz. Li:onard P., and Mu-Ner B. Schaefer 

1936. Descriptions of new intergeneric hybrids be- 
tween certain cyprinid fishes of north- 
western United States. Proceedings of the 
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pp. 1-10. 
Shapovalov, I eo, and Whijam A. Dill 

1950. A check list of the fresh-water and anadro- 
mous fishes of California. California Fish 
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Shapovalov, Leo, William A. Dill, and Almo J. 
Cordon E 
1959. A revised check list of the freshwater and 
anadromous fishes of California. California 
Fish and Game, vol. 45, no. 3, pp. 159- 
180. 
Sharp, Robert P. 

1938. Pleistocene glaciation in the Ruby-East Hum- 
boldt Range, northeastern Nevada. Journal 
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figs. 1-11. 
Shelton, John S. 

1966. Geology illustrated. W. H. Freeman and Co., 
San Francisco, \ii + 434 pp., 382 figs. 
Simpson. J. H. 

1876. Report of explorations across the Great Basin 
of the Territory of Utah for a direct wagon- 
route from Camp Floyd to Genoa, in Car- 
son Valley, in 1859. Engineer Department, 
U. S. Army, Ciovernnient Printing Office, 
Washingt(.)n, 518 pp., illust. (including ap- 
pendices by other authors). 
Smith. Gerai d R. 

1966. Distribution and evolution of the North 
American catostomid fishes of the subgenus 
Pantostciis, genus Calostoinus. Miscella- 
neous Publications of the Museum of 
Zoology — University of Michigan, no. 129, 
132 pp., 22 figs., 1 pi. 
S.MiTH, Gerald R., W. I . Stokes, and K. F. Horn 
1968. Some late Pleistocene fishes of Lake Bonne- 
ville. Copeia. 1964, no. 4, pp. 807-816, 
fiiis. 1-6. 



VOL. VII HUBBS, MILLER, & HUBBS— GREAT BASIN RELICT FISHES 



253 



Snvder, Charles T., George Hardman, and F. F. 
Zdenek 
1964. Pleistocene lakes in the Great Basin. United 
States Geological Survey, Miscellaneous 
Geologic Investigations, map 1-416. 
Snyder. Charles T., and Walter B. Langbein 

1962. The Pleistocene lake in Spring Valley. Nevada, 

and its climatic implications. Journal of 
Geophysical Research, vol. 67, no. 6. pp. 
2385-2394, figs. 1-5. 

Snyder, John Otterbein 

1908. Relationships of the fish fauna of the lakes 
of southeastern Oregon. Bulletin of the 
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doc. no. 636, pp. 69-102, figs. 1-4, 1 map. 
1917. The fishes of the Lahontan system of Nevada 
and northeastern California. Bulletin of the 
United States Bureau of Fisheries, vol. 35, 
for 1915-16, doc. no. 843, pp. 33-86, figs. 
1-9, pis. 3-5. 

Stewart, Norman Hamilton 

1926. Development, growth, and food habits of the 
white sucker, Catostomus commersonii 
LeSueur. Bulletin of the United States 
Bureau of Fisheries, vol. 42, no. 1007, pp. 
147-184, figs. 1-55. 

Stokes, W. L., G. R. Smith, and K. F. Horn 

1964. Fossil fishes from the Stansbury level of Lake 
Bonneville, Utah. Proceedings of the Utah 
Academy of Sciences, Arts and Letters, vol. 
41, pp. 87-88. 

Strahler, a. N. 

1963. The earth sciences. Harper and Row, Inc.. 

New York, vii + 681 pp., many figs. 
Sumner, F. B., and Urless N. Lanham 

1942. Studies of the respiratory metabolism of warm 
and cool spring fishes. Biological Bulletin, 
vol. 82, no. 2. pp. 313-327, figs. 1-4. 
Sumner, F. B., and M. C. Sargent 

1940. Some observations on the physiology of warm 
spring fishes. Ecology, vol. 21, no. 1, pp. 
45-54, figs. 1-2. 
Tanner, Vasco M. 

1950. A new species of Gila from Nevada (Cyp- 
rinidae). Great Basin Naturalist, vol. 10, 
nos. 1-4, pp. 31-36, I fig. 



Thompson and West 

1881. History of Nevada. . . . See entry under Myron 
Angel, Editor. 
Thornbury, William D. 

1969. Principles of geomorphology (Second Edition). 
John Wiley and Sons, New York, xi + 594 
pp., many figs. 
Uyeno, Teruya 

1960. Osteology and phylogeny of the American 

cyprinid fishes allied to the genus Gila. 
Doctoral Dissertation, University of Mich- 
igan, 174 pp., 10 figs., 35 pis. 

1961. Late Cenozoic cyprinid fishes from Idaho with 

notes on other fossil minnows in North 
America. Papers of the Michigan Academy 
of Science, Arts, and Letters, vol. 46, for 
I960, pp. 329-344, figs. 1-3. 
Uyeno, Teruya, and Robert Rush Miller 

1962. Relationships of Empeirichthys erdisi, a Plio- 

cene cyprinodontid fish from California, 
with remarks on the Fundulinae and Cyp- 
rinodontinae. Copeia. 1962, no. 3, pp. 
520-532, figs. 1-7. 

1965. Middle Pliocene cyprinid fishes from the 

Bidahochi formation, Arizona. Copeia, 
1965, no. I, pp. 28-41, figs. 1-8. 
Wheeler, Geo. M. (assisted by D. W. Lockwood) 
1875. Preliminary report upon a reconnaissance 
through southern and southeastern Nevada 
made in 1869. United States Geographical 
and Geological Surveys West of the One 
Hundredth Meridian, vol. 1, pp. 1-72. 
1889. Report upon United States Geographical Sur- 
veys West of the One Hundredth Meridian, 
in charge of Capt. Geo. M. Wheeler, vol. 1, 
Geographical Report, Washington, 780 pp., 
38 pis.. 3 maps. 
Wheeler, Sessions S. 

1971. The Nevada desert. The Caxton Printers, Ltd., 
Caldwell, Idaho, 168 pp., many figs. 
Wilson, Brian L.. James E. Deacon, and W. Glen 
Bradley 

1966. Parasitism in the fishes of the Moapa River, 

Clark County, Nevada. Desert Research In- 
stitute, University of Nevada, Preprint Series 
No. IS, 18 pp., 1 fig. 



SUPPLEMENTARY NOTE 

Frequent reduction in occurrence of barbels in isolated spring populations of Rliinichfliys osculiis ( in southwestern 
Oregon) has been noted by Bisson and Bond (Copeia, 1971. no. 2. pp. 274-275). A general trend may well be 
indicated (see p. 104). 



INDEX 



98 



98 



15, 19- 



47-48, 54, 5(1, h()-6l. 



153 



151, 205, 23 K 234, 239- 



!l. 33, 232 



Acrocheihis, 193 

Adaptations to life in fast water, 98 

Adaptations to life in springs, 76. 81-82, 84, l()()-l()4, 

119. 124-127. 132-133, 136, 145, 178. 181 
^.i,'<'.v/«. 97, 193-194 

chrysoi^'cister. 93 

rohiisla. 104 
Afiosia ( Apocope 
' adobe, 98 

coiiesii. 98 

fahala, 98 

nevculeiisis 

niihila. 98 

oseiihi. 98 

uiiHitillii, 98 

velifem, 98 

V(//7<Mr/, 98 
Antelope Valley, 

1 14. 233 
Apocope. 97-98. 104 

osciila niihilci. 98 
oscidii. 98 
Archeological sites, fish in 
ArcliopUtcs inlcrruplus. 58 

240. 244 
Area and diversity, 77-19 
"Area of Sterile Basins." 15. 
Argyreiis nolahilis, 98 

niihiliis, 98 

osciiliis. 98 
Ash Meadow. 77 
Ash Spring. 232 
Azieciila villcita. 93 
Bass — see Micropwrus sulnioides 
Bassett Lake, 239-240 
Behnke, Robert J.. 206. 238. 244 
Berwick, Stephen H., 135, 244 
Big Shipley Spring, 17-18, 76, 99. \W. 113. 115. 

117-121. 125. 127. 154. 230, 242 
Big Smoky Valley. 10. 14-15. 72. 123-125, 238 
Big Sprinus, 23 1 

Bifateral asymmetry. 84, 89-90, 93-94, 134. 
Birch Creek, 117-119 
Birch Ranch Springs, 99, 115. 118-121. 124, 

154-156, 172. 242-243 
Bishop Creek. 99. 104. 106-107, 109, 113, 

120, 123-124, 126, 148, 151-152, 
168-172. 175. 178-180. 242 

Bluegills — see Lcpoiuis nnu rocliinis 

Boone Springs. 54 

Box Sprinus, 14. 20 

Brown. Robert E.. 20. 1 14. 244-245 

Brues. C. T.. and wife. 228 

Bull Creek, 33 

"Bullheads" — sec Iciidiinis (Ameiiini.s) species 

Butte Creek, 44, 201 

"Butte Lake," 45 

Butte River (pluvial), 39, 44-46 

Butte Valley. 39. 44^6, 55, 58, 176. LSI. 192. 196. 

199-202. 206. 217-219. 223. 226. 235. 243 
Cain. Donald R., 205-206, 244 
Callaghan Ranch spring, 123 



149, 


, 190 


127, 


147, 


1 16, 


1 19- 


155, 


158, 



Callaghan (Woodward) Creek, 14, 121 

Campbell. Mr.. 204-205. 245 

Canyon springs. 3. 75. 96, 158 

Carassius aiinitiis. 18. 58. 115. 157, 204-207. 239, 242 

Carico Lake Valley. 10. 87. 90. 99. 104. 106, 109, 113, 

120, 125-126. 128. 133. 241. 245 
Carp — see Cvpiiniis carpio 
Carp X goldfish hybrids. 58. 204 
Carpenter River (pluvial). 61. 64-66 
Carson River. 143. 145. 148. 151-153, 155. 172 
"Catfish" — sec Icfcdiinis ( Anieiiinis) species 
Catostoiuiis {Catostoiuiis) . 74 
coiunwisoiiiiii. 1 36 
tahocnsis, 74. I 5 I 
Calostoniiis (Pcintosteus) , 74. 230 

platvr/ixncliiis, 64. 74. 151. 229-230. 233. 241 
Cave Creek. 44-45. 198-199. 226 
Cave Valley. 55. 65 
Cavender. Ted M.. 210. 245 
Ceraloplirlhiiii. 134 
Cham. 29. 107. 114. 121. 129. 157. 168. 199. 201-204, 

206-207. 225-226 
Characters of fishes. 79-96 
Charnock Springs. 15 
Chase Springs. 32 
Clui.sDiistes ciijiis. 74 
Cherry Creek. 206. 234 
Cherry Creek Hot Springs. 206 
Chromo.somes, 193, 195-196 
"Chubs" — see Gila hicolor 
Cleve Creek. 64, 234, 237 
Clinostomus cloni;aliis. 94 

Clover Valley, 30-32, 76, 99, 126. 129-134. 175. 242 
Clover Valley springs. 129-134. 136 
Coffin. Patrick. 106. 245 
Coils Oeek. 19. 76. 8(1. 99. 104. 113-116. 118-121. 

124. 127. 242 
Cold Creek. 26. 157 
Cold Spring. 19, 157 
Colorado River. 32-39. 47. 55. 62. 64-68. 71. 73-75, 

97-99, 105, 230, 24(1 
Columbia River, 1. 7, 75. 97-99 
Comins ( ="Cummings" or "Cumin") Lake (also 

Meadow and Spring). 59-60. 231 
Corei^diuis <■. chipeaforniis, 238 
Cottits hcldini;i. 74 
Coiicsiiis. 210 

pluiuheus. 210 
Crane. Forest Ranger. 19. 20. 232. 245 
Crappie, black — see Pomoxis nii^ronuuulaiiis 
Crenicluhvs. 36. 73. 75. 227 

hailcri. }>. 71. 227-228, 241, 244 
ncradae. 7. 35-36, 71-73, 75-77, 227-229, 235. 244 
Crescent Valley, 10-11. 15. 75. 82, 99, 104, 106, 109, 

I 13. 1 18-121. 123-124. 126. 128. 241 
Crouch, David Cr., 246 
Crystal Spring. 232 
Currant Creek, 33, 228 
Crpriniix carpio. 58. 66. 134. 151. 157. 202. 204-205. 

234-235. 238-239. 242 
Davis. Charles M.. 246 



VOL. VII 



INDEX 



255 



129. 131. 201 

35. 74, 77. 



204-205. 207. 228. 



1 48 



244-245 



88. 109, 113-121 
146-147, 153-156 
180. 230, 239. 243 
76, 80. 90, 114. 118 



209 



244 



Deacon. James E 

240. 245 
Death Valley system 
Deep Creek. 61 
Deep Creek. West. 105 
Delamar Valley, 65 
Desert Fishes Council. 
Devils Gate. 16-17 
Diamond Springs. 18 
Diamond Valley, 15-19, 76 

125, 132-133, 136. 

161, 172. 173, 177. 1 
Dianas Punch Bowl, 19-21 

127. 242, 244 
Dice, Lee R., 245 
Dixie Valley, 72 
Dodge, Frank N., 205-206 
Dry Lake. 21 
Dry Lake Valley. 65 
Duck Creek. 57. 205 

Duck (=Lake) Vallev. 55. 63. 65-66. 23 
Duckwater Creek, 33. 36, 77, 142. 228 
Duckwater Spring. 227-228 
Duckwater Valley. 228 
Dwarfism, 136, 140, 155. 177, 
Eagle Lake, 77, 144-145 
Egan Creek, 58-59, 235, 245 
Egan Creek Valley, 57-59, 77 
Egan Valley. 5 
Emerald Lake and Cave. I 8 
Emigrant Valley. 36 
Eiiipenichthvs, 73. 227, 240 

lato.s- latos, 229. 235 
Endemism. 70-76. 99. 121-141, 150, 153 
Ecjiiiscliini. 1 14 
Eremichlhvs. 181, 193-194 

acros. 81, 193 
Extinction, 93, 123, 236, 240, 242 
Faunal depauperation. 70-79, 95-96, 123, 

204, 236, 240-244 
Ferguson Creek, 19 
Fife", Otto, 70, 245 
Fish Creek, 22, 25. 169. 240 

Fish Creek Springs, 23, 25-26. 80, 157, 159, .. 
Fish Creek ( Little Smoky ) Valley, 22, 25-26, 38 

160, 168-174 
Fish Lake, 36, 79 
Fish Pond Springs, 60, 205-206 
Flower (=Flowery) Lake, 54, 202-203 
Flynn Pond, 17, 239 
Follett, W. I., 245 
Fossil fishes, 73 
Franklin Lake, 44, 198 
Franklin River (pluvial), 43-45, 198 
Fundiihis, 227 

parvipinnis, 227 
Gambiisiti affinis. 236. 241 
Gardner Creek. 198 
Gardner, W. J., 198, 245 
Gasterosteiis aculcatiis, 144 
Gila, 94, 183-191, 193-194, 2 
Gila (Gila), 74, 142, 182, 193 
atraria, 58, 60, 64, 66, 142- 

234, 241, 244 
copci, 93, 189 



124- 
158- 

-121, 



-181 



197-199, 



168- 



174 
57- 



14, 



143, 193, 206, 226, 229- 



jordani, 232 
orciitlii, 214 
rohiista, 1 96 

jorduni, 229. 232 
Gila (Siphalele.s), 15. 123, 182, 190, 193, 209 

bicolor, 1, 13, 18, 23, 25, 35-36, 38, 41, 51, 71-77, 
79, 81-94, 121, 135, 142-180. 182. 187, 189, 
193, 196, 198, 208-209. 212, 214, 221-223, 
228-229, 241-242. 245 
bicolor, 150 

eiichila, 25-26, 80, 89, 142, 145, 147-148, 150. 

152-155, 157, 159, 160-174, 177-180, 243 

isolala. 31-32, 80, 89, 91, 134, 142, 145, 147-148, 

150, 152, 154-167, 170-180, 183-191, 236, 

242-245 

inohavensis, 1 46 

newarkensis, 26, 86, 89, 142, 145-148. 150, 152- 

174, 177-180, 243 
obesa, 18, 23, 26, 71-72, 76, 87, 89, 93-95, 115, 
142-156. 158. 160. 161-175. 177-180. 192. 223. 
242-243 
peclinifcr. 89. 93, 95, 142-151, 153. 242 
subspecies. 232 
nbc.ui. 77. 143 
Gila River. 97 

Goldfish — see Carassius iiiiratiis 
Goose Lake drainage hasin. 77 
Goshute Creek, 237 
Goshute Lake, 48, 55-58. 234 
Goshute Valley. 47-48, 54-55, 57, 181, 196. 202- 

208. 213, 219, 243 
Grass Springs. 58. 205 
Grass Valley, 10-15. 82. 106-107 

238. 242, 245 
Grass Valley Creek, 121-128 
Grass Valley Spring, 80, 99 
Grayling — see I Itynuillii.s sii;iulcr tricotoi 
Green River, 97 
Greenbank, John T., 91, 245 
Gxraiiliis, 28 
Hall Creek, 106 
Hardy Creek. 203 
Hcsperoleuciis, 193 
Hippums, 204, 207 
Horse Lake drainage hasin. 77 
Hot Creek. 33 

Hot Creek Lake (ancient). 5. 33-34. 77 
Hot Creek Valley, 22. 33-34. 36. 38. 232 
Hot Spring (Antelope Vallev). 20 
Hot Springs ( Ruhv Valley), 199 
Hot Springs Valley, 142 
Hubbs, Clark, 228 

Humboldt River, 10, 15. 22-23, 25, 
39, 72-74, 80, 99-100, 104, 
118-121, 124, 126, 
151-153, 155-156. 



204. 
109. 121-128, 237- 



27, 30-31, 35, 
106-113, 115- 
131-132, 143, 145- 
58, 160, 161. 168- 



175, 177-179, 192. 223. 230 
Huntington Creek. 22-23. 27. 169 
Huntington Vallev. 25 
Hatchings, Selar .S.. 70, 245 
Hybopsis, 210-211 
craineri, 244 

(Eriiuysta.x) disyimilis. 21 I 
( Eriiuysta.x ) .x-puiictaliis. 2 1 1 



-231. 237-239, 



38- 
116. 
148, 
172. 
242 



256 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Hybrids. Ciiloslomiis platyrlniu hiis icihocnsis, 151 

Carp ■ goldfish. 58. 204 
htahinis (Aiiiciunis) species. 17. 157. 239 
nwlas ijicliis. 151. 239 
iichiilosiis, 239 

nchiilosiis. 151. 239 
lllipah Creek. 68, 237 
Illipah Sprina. 68 
Independence Valley. 30-32, 48, 76. 99. 122. 1 2h, 134- 

140. 147. 172. 175-180. 236. 242-243 
Indian Creek, 75, 80. 99. 106-107. 123-124. 192. 241 
Indian Ranch Spring, 14 
Indian Spring, 206 
Introductions of fishes, 17-19. 31, 45-46, 55, 58, 60, 

64, 66, 68, 70, 115, 123, 135, 151. 157, 198, 

205, 207-208, 227, 229. 231. 235-240. 245 
Iowa Creek. 106 
Ireland, W. E., 207 
Jacobsen, Jorse, 115, 245 
Jakes Valley, 29, 55, 67-68, 237 
Jakes Wash, 67-68 
Johnson Ranch springs, 54, 202-203 
Josephine Spring. 17 
Jitiuiis. 229 

Karyotype, 193, I95-I9fi 
Kingston Creek, 238 
Klamath Lake, 150 

Knutson, Mrs. (formerly Mrs. Dick McGee). 123 
Kobeh Valley, 10, 15-16, 19, 114 
Lackawanna Spring, 206 
Lackey, Martha B.'^ 246 

Lake Antelope, 6-7, 38, 47-49, 54-55, 60-62, 70-71 
Lake Bonneyille, 1, 7, 9. 29. 47. 54. 56, 60-63, 65-66, 

68-71, 73-75. 77. 79. 99. 105, 143, 230-231. 

235, 237 
Lake Bristol, 64-65 
LakeCarico, 10, 99 

Lake Carpenter, 5-7. 47. 55. 62-6h. 71. 74. 231 
Lake Cave. 47. 55, 65 
Lake Clover. 5-7, 10. 28-32. 39. 42-43, 47. 71-74. 76. 

99-100. 106. 126, 129-141). 145. 160. 175-180. 

238 
Lake Coal. 33. 64-65 
Lake Columbus, 74, 78 
Lake Crescent, 10, 99 
Lake Delamar, 64-65 
Lake Diamond. 6-7. 10. 15-23. 26, 33, 71-74, 76, 99- 

100, 104-106, 113-121, 127, 135, 145. 153-156, 

238, 242 
Lake Diana, 6-7, 10, 15-16, 20-21. 71. 76. 99 
Lake Dixie. 18, 74, 142 
"Lake Dou-pah-gate," 17. 239 
Lake Franklin, 5-7. 22, 26-27. 29-31, 38-48. 54-55. 

58, 70-71, 73. 76-77. 99, |(1X. 150. 196-199, 

215-224. 238. 243 
Lake Gale, 6-7, 26-27, 38-39. 42. 45-47, 54-55. 58. 

67. 70-71. 73. 77. 99. 181. I9(i. 199. 201-202. 

217. 243 
I ake Ciilbert, 5-7, 10-16. 71-74. 76-77, 99. 106. 121- 

128. 238. 242 
Lake Ciroom. 36 
Lake Huhbs. 6-7. 10. 22-24. 26-29. 39. 42. 45. (i4. 67. 

71 
Lake Jake. 5-7. 22. 26-27. 33, 45, 47, 55, 64-68, 7 1 , 74 
Lake Kawich, 35 



Lake Lahontan, 1, 7, 10. 23. 27. 29. 32-38, 42-43, 64, 

71, 73-75, 79, 104, 142-151, 169, 175, 231 
Lake LeConte, 57 

Lake Lunar Crater, 6-7, 22, 32-33, 35-38, 71 
Lake Manly, 74, 79, 235 
Lake Mead, 239 
Lake Newark, 5-7, 10, 15. 21-27. 33. 35, 37-39, 64, 

67, 71-74, 76-77. 99. 145. 147. 156-174. 177- 

178 
Lake Penoyer. 35 

Lake Pine '( =Lake Wah Wah ). 6-7. 68-70 
Lake Railroad. 6-7. 15. 21-22, 25, 32-38, 65, 67, 71- 

78, 142, 146, 227-229, 232 
Lake Reveille, 34, 78 

Lake Snyder, 6-7, 16, 22-23, 32-33, 35, 37-38, 71 
Lake Spring, 6-7, 9. 47. 55. 60. 61-66. 71, 196, 207- 

208, 229-231 
Lake Steptoe, 6-7. 38. 45. 47-49. 54-60. 62, 65. 67, 

70-71, 73, 76-77, 99, 181, 196, 204-207. 217, 

231, 243 
Lake Steptoe. Lower. 55. 59 

Lake .Steptoe. Upper. 6-7. 38. 57-60. 70-71. 204. 231 
LakeTahoe, 144 

Lake Toivahe. 10. 14-15. 21. 72. 74, 99, 121. 142. 238 
Lake (=buckl Valley, 55. 63. 65-66. 231 
Lake "Wah Wah" (:=Lake Pine). 68-70 
Lake Warini;. 6-7, 29, 38-39, 42-58, 60-62, 70-71, 

73, 76-77, 99, 196-197, 202, 204, 215-224, 243 
Lake White Mountains, 79, 142 
Lake Winnenuicca, 147 
Lake Yahoo, 6-7, 10. 15-16. 20-22, 71, 76 
La Rivers, Ira, 123. 245 
Lehman Creek, 237 
Lenina, 207 

Lepoinis nuu iiu /lii us iiuunn liirus. 239 
Leptocolliis (iniuiliis. 90 
Leiicidius. 144 

pi'dinifcr. 143, 147. 148 
Lewis. Donald H.. 107. 135. 198-199. 245 
Little Fish Lake. 142 
Little Fish Lake Valle\. 33. 36 
Little Fish Lake(s). 3. 6-7. 21-22. 25. 32-33. 36. 71- 

73. 76 
Little Meadow Lake. 240 
Little Round Valley ("Mosquito Flat"). 37 
Little Smoky Valley (=Fish Creek Valley). 22. 25-26, 

38, 157, 16,8-174, 240 
Little Soda Lake, 148 
Lockard, Dale V.. 205. 232. 235. 245 
Lockes Ranch springs. 228 
Long Valley, 26-29^77 
Long Valley Slough, 29 
Lucunia purva, 108, 221 
Lugaski, Thomas P., 123, 206, 245 
Malheur Lake drainage basin, 77 
Mangum, Earl, 205-206, 240, 245 
Manse Spring, 235 
Maps consulted, 8-9 
Miiii,'(iii.sciis. 210 
Martonyi, Louis P., 246 
Mayuard Lake, 65 
Mcbermitt Creek. 54. 57. 204 
McGee. Dick. 121, 123 
McGill. William. 206. 245 
Meristic correlations, 88-89, 93 



VOL. VII 



INDEX 



257 



Meristics. temperature effects. 84-85, 90-91 
Micropterits sahnoides, 45, 60, 77, 109, 135, 151, 198- 
199, 204, 206, 230, 234, 242-243 

sahnoides. 239 
Miller, Francis Hiihbs, 246 
Minnie Spring, 198-199 
Miocene. 3 
Moapa. 181. 193-194 

coriacea. 193. 229. 232 
Moapa River. 232 
Mohave River. 146 
Monitor River (pluvial). 16, 20. 99 
Monitor Valley. 10, 15-16, 19-21. 104. 113-115, 237 
Monte Neva Hot Springs, 58, 204 
Moore, Billy. 245 
Moorman, Mr., 68, 245 
Moriarty. James R., 246 
"Mosquito Flat" (Little Round Valley), 37 
Mound Spring, 32 
Murphy Spring, 231 
Murrav (=Murry) Creek, 57-58, 205 
Myiiophylliim. 134, 151. 201 
Ndids marina, 1 15 
Nasturtium. 106, 121, 129. 157. 198-199. 201-206. 

225-226 
National Science Foimdation. 246 
Nelson Creek, 43. 54, 202-203 

Newark Valley. 22-26. 33. 35. 146, 156-169, 239, 243 
Newark Valley springs, 157 
NocoDus icptocephalus, 93 
Notropis saladonis. 93 
Odgers Creek. 46. 201 
Oregonichthys, 210, 245 

crameri. 210, 245 
Orthodon, 94, 193 
Osteology, 182-191 
Owens Valley, 238 
Pahranagat Valley, 36, 65, 232 
Pahrump Valley, 235 
Paiitosteus. 230 

dclphinus. 230 

jonlaui. 230 

lahonlan. 230 
Paris Creek, 46 
Parker, Elizabeth, 246 
Patterson Wash, 66 
Penoyer (Sand Spring) Valley. 35 
Perca flavesceiis. 151. 239 

"Perch" — see Archopliles intcrniptus. Perca flavcscens 
Phalan Creek. 54. 57. 202-203 
Phoxinus. 94 

lagowskii variegatu.s. 94 

percnurus, 94 

pho.xiinis, 94 
Pine Creek. 15. 237 
Pine Grove Creek. 70 
Pine Valley, 69 
PisidiuDi, 28 
Plaaopterini, 74 
Pluvial lakes, 10-70 
Pole Creek, 43-45 
Poiiw.xis nigromaculatus, 240 
Pony Springs, 66 
Population "structure, 96. 113. 121. 128. 134. 140. 223- 



Potamogeton. 134. 151. 168. 203. 226 

cf. P. pcctinatus, 107. 113-114, 129, 151, 168, 201- 
207. 226 
Potts, George, 19-20, I 14, 245 
Potts Ranch Hot Springs, 19-20, 80, 99, 113-114, 118- 

121. 127. 242 
Procamharus ciarkii. 236 
Prosopium. 73-74 

wii/iamsoui, 74 
Pyramid Lake. 143-144 
Railroad Canvon. 16-18 

Railroad Valley. 7, 32-38, 65. 142, 227-229. 240. 244 
Ralph. Mr., 129 
Ralph's Warm Springs, 134 
Ralston Creek, 19 
Ramirez Springs, 199 

Rami catesheiuna. 131, 135, 226, 236, 241-242 
Reese River. 10. 99. 192. 238 
Refugia for fishes, 232, 235, 241 
Relictus. 180-196 

solitariiis. 7, 31, 41, 43-46. 48. 51. 53-55. 58. 61), 64. 
70-71. 73, 75-77. 7^>-92, 94-95. 98-99. 107. 
123-124. 131. 175-176, 180-226, 229-230, 
232-235. 240-241. 243-245 
Reveille Lake and Valley, 34 

Rhinichthvs. 91-9H. 100. 123. 125. 133. 136. lSl-191. 
193_194, 196, 208-212, 214, 222 
atratidus. 97-98, 1 82 
calaractac, 97-98, 1 82 
i'vcrmamu, 91-9H 
falcatus, 97-98, 182, 193 
lariversi, 15. 72, 123 
nubilis. 98 

rohuslus. 105 
osculus, 1. II. 13. 15. 18-21. 23. 41, 51. 70-77. 79- 
92. 94. ^6-141, 151. 154. 182, 193. 208. 221. 
223. 241 
adohc. 105 
lariversi, 123-125 
k'thoporus, 31-32. 80. 91. 99-104. 110-112. 118. 

122. 124-129. 131-141. 175. 236. 242-243 
nevadensis, 94 

nuhilus. 105 

oligoporus. 31. 76. 99. 100-104. 110-112. 118. 

124-141. 175, 242 
reliquiis. 14, 80, 86, 99-104, 106, 109-112, 118, 
121-128. 132-133, 135-141, 182-191, 240, 
242, 245 
robustus. 19, 71-72. 76. 80. 86-87, 95, 97, 99-121, 
123-128, 131-133, 135-141, 151, 175, 192, 
199, 222-223, 229-230, 241-245 
thcrmalis. 97 
xarrowi, 105 
nmatiila. 97-98 
Richardsonius egregius. 74, 108-109, 151, 229-231, 241 
Roberts Creek, 19, 99, 113, 115, 238, 245 
Robinson Creek and Lake, 44 
Rohison, Bert, 64. 207. 234-235. 245 
Rowe. Phil. 208 
Ruby Lake. 44-45. 77. 108. 150. 198-199. 225. 230- 

23 1 . 243 
Rubv Valley. 39, 80, 99-100, 107-109, 113, 119-120, 
126, 181, 196-199, 209, 217-219, 225. 235. 243, 
245 



258 



CALIFORNIA ACADEMY OF SCIENCES 



MEMOIRS 



Russell River (pluvial), 33 

Ruth Pond. 205 

Sacramento perch — see Archoplitcs iiucniiptus 

Sacramento River. 104 

Sahno. H). 45. 60. 6fi. 70. 74-75. 99. 121. 202. 20(-i. 236 

tii;iitih(iiiilti. 238 

clarkii. 64, 73-74. 236-237. 243 
hcnshawi, 237-238 
lewisi. 237-238 
iiloh. 237 

miidncrii. 18, 55. 58. 123. 129. 131. 169. 204. 234- 
235, 238 

tntftci. 237 
S,ilvcliiiii\. 202 

loulinalis. 55. 58. 123. 169. 204. 238 
Salver. J. Clark. 198 
Sand .Spring Valley. 36-38 
.Sangamon (post). 3 
•Sara, Isador, 25, 169. 245 
■Sawmill Creek. 59 
.Scale structure. 192-193. 214 
.Schell ( =Shell ) Creek, 207, 234 
Schell Creek Spring, Lower and Upper, 207 
Schell (=Shell) Creek Vallev. 233-234 
Schell Vallev, 234 
Schultz, Robert I.., 207 
Scirpiis. 207. 229 
Seiiiotiliis iilroinacithitiis. 94 

iiuirgcirita, 210 
Sevier Lake. 69 
Sevier River, 230 
.Sexual dimorphism, 84. 113. 120. 125, 128, 133. 136, 

140, 152, 15h, 161, 166. 172. 179. 219-222 
Shatter Valley. 47, 55 
Sheep Corral Springs, 201 
Shell (=Schell) Creek and V.illey, 233 
Shipley Hot Spring, 17 
Shirlev, Howard G., 246 
Shor, Llizabcth N., 246 
Shoshone relugiuni. 232. 235 
Shoshone Springs, 64, 23 1, 234-235 
Siphatcles, 15. 123. 142-144. 148 

hicokir, 1 44 

ohcsiis. 94. 142, 144, 147 

pccrinilcr. 1 44 
Skull Creek, 14. 237 
Snake Valley, 63, 87, 23 1, 233, 237 
Snow Creek, 46. 201 
Snow Water Lake. 31. 131 
Snyder. C. T.. 23. 39. 245 
Snyilcrichlhy.', {—Gilti) copci. 93 
Sodaville hot springs. 229 
Soldier Meadows. 97 
Soulages. Robert H., 19. 245 
Spring (Vallev) Creek. 64. 207, 230, 233-235 
Sprin^ Vallev. 21. 47. 62-64. 196. 207-208. 217. 219. 

223. 226, 229-235, 237, 240-241 
Spruce Point Spring, 32 
Scpialius (ilicicic. 93 
Steiner Creek, 14 

Steptoe Creek, 57-60, 205-206. 232. 234-235 
Steptoe River (pluvial). 38. 47. 56-58 
.Steptoe Vallev. 29. 47. 54-60. 181. 1 9h, 202. 204-207. 
219. 225. 233-235. 237. 239-240. 243 



Stone Cabin Creek. 19 

Stone Cabm ( =Willow Creek) Valley, 142,232 

.Stoneberger Creek, 19-20. 99. 113-1 14, 238 

Stratton, Jerome Phalan, 29. 46. 199, 201. 203. 206. 245 

Slypoilon si};nifcr. 93 

Slicker — see Catoslonuis {I'aiUostcits) plalvrhxncliiis 

Sulphur Spring, 18, 76, 82, 146-147, 154-156,' 158, 160, 

161, 166, 168, 172, 179, 234, 243 
Survival of endanuered species and other local forms. 
115. 131. 134-135, 154-160, 175-180, 197-199. 
202, 206-207, 227-229. 239-242 
Tea Creek. 108 

Temperature effect on meristics, 84-86, 90-91, 93 
Temperature resistance, 1 14 
Thirty-mile Spring, 206 
ThyimiUus signifcr iricuhn . 238 
T'uiroi;u. 193 

Tickaboo ( Desert ) Valley, 36 
Tippett Valley, 60 

Town Creek. 99, 104. 106-108, I 19-120, 124 
Town Creek, sprinijs near, 109, 1 13, 123, 126 
Transferences of fishes, 58, 60, 64. 66. 107-109. 207- 

208. 229-235 
Trelease. Thomas J.. 135. 229, 231, 245 
Trophic adaptations, 92, 136, 143. 146-150, 153 
Trout — see Salnio, Scilvc'linu.\ foiitiinilis 

Brook — see Scilreliiius fonliinilis 

Ciolden — see Scilmo ugitabonila 

Lahontan cutthroat — see Saliiio cUirkii hciisluiwi 

Rainbow — see Salnio gairdiierii 

Yellowstone CLitthroat — sec Salnio clarkii Icwi.si 
Trout Creek, 237 
Truckce River. 77 
Tule Dam Spring, 18 

Twin Springs. 19. 36. 46, 54, 142. 201. 203, 232 
Umpqua River, 97 
Utah Lake, 69 

Uniciilaiia. I 15, 168-169, 201, 203. 226 
Uyeno. Teruva. 142. 193. 205, 245 
Vallee, Ellen J., 205, 245 
Vauuhn. James M., 206 

Wah Wah Drv Lake, Springs, and Valley, 69 
Walker, BovdW.. 232 
Walker Lake, 144 
Walti Hot Springs. 11.14 
Waring Valle\, 235 
Warm Spring ( Monitor Valles), 19 
Warm Spring Valley, 228 

Warm Springs ( i- Clover Spring ), 31, 99, 126, 129-134 
Warm Sprinas (Independence Vallev), 31-32, 76, 80, 

91. 99, 131, 134-140, 171. 175-180. 236 
Warm Springs (Newark Valley). 157. 239 
Warm .Springs Pond ( =Warm Springs). 239 
Weslwood. Vernon. 32. 245 
While Lake. 62, 64 

White Mountains (Fish) Lake. 79. 142 
White River (pluvial). 33. 35-36. 47. 55. 62, 64-66. 68, 

"'"'7— 2 ""9 2^2 
White River Valley, 240 

Whitefish, Great Lakes — sec Coicgoitus c. cliipcafonnis 
Williams Creek, 60, 204 
Williams River, 97 
Willow Creek, 58, 60. 204 

Willow Creek (=StoneCahm) Valley. 142,232 
Wilson Creek, 66 



VOL. VII 



INDEX 



259 



Wisconsin (late), 3, 27 
Worthington Spring, 63-64, 235 
Wrights Spring, 201 
Xxraiichcn. 74 
Yahoe (Yahoo) Creek, 21 



Yelland, John, 204-205, 207, 2^1, 234-235, 237 ''40 

245 
Yoakum. James, 232, 235. 237. 245 
Zahuranec, B. J., 228 
Zubiri, Mrs., 203. 206, 245 



7X 



The Origin of Birds 

AND THE Evolution of Flight 



Edited by Kevin Padian 




■f 



■\\ 



"-'?, 



KvMt/ER 
Ohprnr lura Firhstatf.Pavei'n 




Published by 

California Academy of Sciences 

San Francisco 

1986 



Memoirs of the California Academy of Sciences Number 8 



•'v/l/\. 



w'ES 



The Origin of Birds and the Evolution of Flight 



The Origin of Birds and the Evolution of Right 



Edited by 
Kevin Padian 



Published by 
California Academy of Sciences 



^mmmm 



^^^ 




San Francisco 
1986 



Memoirs of the California Academy of Sciences Number 8 



© 1986 by the California Academy of Sciences, Golden Gate Park, San Francisco, CA 941 18 
ISBN 0-940228-14-9 



Contents 



Preface — Kevin Padian. 



Saurischian Monophyly and the Origin of Birds— Jacques Gauthier 

Introduction _ 1 

The Thecodont-Bird Hypothesis 1 

The Mammal-Bird Hypothesis 2 

The Crocodile-Bird Hypothesis 3 

The Dinosaur Hypothesis 4 

Materials and Methods 7 

Introduction to the Basic Taxa 8 

Elmisauridae* Osmolska, 1981 9 

Caenagnathidae Sternberg, 1940 9 

Ceratosauria Marsh, l&&4b (n. comb.) 9 

Camosauria Huene, 1920 (n. comb.) 9 

Omithomimidae Marsh, 1890 (n. comb.: includes Deinocheiridae of Osmolska and Roniewicz, 1970) 10 

Deinonychosauria Colbert and Russell, 1 969 1 1 

Troodontidae Gilmore, 1924 (=Saurornithoididae Barsbold, 1974) 11 

Dromaeosaundae Matthew and Brown, 1 922 1 1 

Avialae (n. txn.) 1 1 

Omithurae Haeckel, 1866 13 

Aves Linne, 1758 14 

Phylogenetic Analysis 14 

I. Phylogenetic Relationships within Dinosauria 1 4 

II. Phylogenetic Relationships within Theropoda 1 8 

III. Phylogenetic Relationships within Tetanurae 23 

IV. Phylogenetic Relationships within Coelurosauria 26 

V. Phylogenetic Relationships within Maniraptora 30 

Conclusions 35 

I. Summary of the Main Phylogenetic Conclusions of this Work 35 

II. The Problem of the Definition of the Name Aves 36 

III. Implications for the Origin of Flight 37 

Acknowledgments 37 

Literature Cited 37 

Appendix A. Archosaur Phylogeny: On the Relationships of Certain Extinct Archosaurs to Extant Crocodiles 

and Birds 42 

Appendix B. Taxon/Character Matrix for Saurischia Data Set 47 

Addendum 47 



The Arboreal Origin of Avian Flight— Walter J. Bock 

Introduction 57 

The Arboreal Theory 58 

Avian Flight Features 59 

I. Homoiothermy and Feathers 59 

II. Body Size 62 

III. Three-Dimensional Orientation 62 

IV. Bipedalism and the Hindlimb 62 

V. Trunk Shape and Structure 63 

VI. Wing and Tail Structure 63 

Evolution of Avian Flight 65 

Comparison of Terrestrial and Arboreal Theories of the Origin of Avian Flight 66 

I. Pseudophylogenies and Analogous Forms 67 

II. Terrestrial Theories for Avian Flight 67 

III. Arboreal Theories for Avian Flight 69 

IV. The Archaeopteryx Stage 70 

Conclusions 71 

Acknowledgments 71 

Literature Cited 71 



The Cursorial Origin of Avian Flight— John H. Ostrom 

Introduction 73 

The Arboreal Theory 73 

The Cursorial Theory 73 

The Evidence 74 

Discussion 80 

Conclusions 81 

Literature Cited 81 

Mechanical Constraints on the Evolution of Flight— C. J. Pennycuick 

Introduction 83 

Power Required to Fly 83 

Power Margin and Body Mass 83 

Upper Limit of Mass 84 

Frequency Scope 85 

Lower Mass Limits 85 

Body Mass and Potential Diversity 86 

Lack of Diversity in Large Flying Animals 86 

Arboreal Versus Cursorial Origin 86 

Gliding Route to Powered Flight 86 

Relationship of Gliding to Powered Right 86 

Arboreal Gliding as the Basis of Foraging 87 

Pressure for Elongation of the Wing 88 

Pressure for Oscillation of the Wing 88 

Origin from Quadrupedal Gliders— Bats and Pterosaurs 88 

Mechanics of the Bat Wing 88 

Mechanics of the Pterosaur Wing 89 

Ancestral Forms 90 

Origin from Bipedal Gliders — Birds 9 1 

Featherless Gliding Precursor of Archaeopteryx 91 

Evolution of Feathers 92 

Further Adaptations of Flying Vertebrates 92 

The Dual Locomotor System of Birds 92 

The Bird Synsacrum — the Pelvic Lever 94 

Ventilation 94 

Thermoregulation 94 

Evaporative Cooling and the Bird Carina 95 

Cursorial to Arboreal Sequence 96 

Divided Ventricle Indicative of a Tall Ancestor _ 96 

Ancestr> of Birds and Pterosaurs 97 

Acknowledgments 97 

Literature Cited 97 



Preface 



On June 1 2, 1 984 a symposium convened by Dr. Luis Baptista 
and sponsored by the Pacific Division of the American Asso- 
ciation for the Advancement of Science took place at the Cal- 
ifornia Academy of Sciences in San Francisco. The subject of 
the symposium was the origin of flight in birds, a topic that has 
garnered considerable attention in the past decade and is still 
unresolved. Did the ancestors of birds live in trees or on the 
ground? What kinds of animals were they? What kinds of eco- 
logical and aerodynamic pressures led them to develop wings, 
spread them, and go aloft? How did they take off and land? 
What lines of evidence are important or even germane to the 
problem? The authors of the papers in this volume present their 
readers with a spectrum of considered opinion and experience. 
Each author treats the problem thoroughly and reasonably. Why 
then should there be so little agreement among them? 

The lack of consensus on how birds learned to fly is hardly 
surprising, inasmuch as no one was present to witness the pro- 
cess 150 million years ago, and the experiment is hardly re- 
peatable. What possible interest would any legitimate scientist 
have in an issue that is never likely to be proven one way or 
another? To answer that question is to delve far beyond the 
issue of bird flight, into the heart of what makes scientists tick. 
No inference is free of theory; no fact stands alone on a pedestal, 
immune from charges of interpretive bias. History occurs only 
once; many possible roads could have been taken, but only one 
was actually followed. We will never know exactly how birds 
began to fly, but that is only part of the issue. More at stake is 
the issue of how questions like this one should be approached. 

In perusing this volume, an outside observer should keep in 
mind that what scientists think is often less interesting than why 
scientists think what they do. The approaches that these authors 
bring to their topic directly reflect their interests and training; 
it is not simply a matter of the available evidence. To a pale- 
ontologist, the evidence for the origin of flight in birds might 
lie largely within the available fossil record, despite its legendary 
incompleteness and its inability to answer many questions about 
the once-living animals now buried in the rocks. A biologist 
might be justifiably more self-satisfied with the potential of the 
living biota to answer such questions; but not all animals that 
once lived are now alive, and some questions must go without 
answers. 

The present volume underscores the diversity of the evidence 
bearing on the question of the evolution of flight. Jacques Gau- 
thier concentrates on the phylogenetic evidence that places the 
origin of birds squarely within the small carnivorous dinosaurs 
(Theropoda; Coelurosauria) of the Mesozoic Era. His cladistic 
approach, while perhaps not fully endorsed by all taxonomists, 
has the advantage of being explicit, hierarchical, and easily ame- 
nable to further testing. Using sets of shared derived characters 
successively nested within each other, Gauthier shows that birds 
are members, in turn, of the coelurosaurs, theropods, dinosaurs, 
and archosaurs (to mention only a few of the more familiar 
subdivisions of the evolutionary hierarchy). The nested se- 
quence of shared derived characters shows us not only the re- 
lationships of birds, but the sequence by which they acquired 
those characteristics commonly associated with flight. It may 
be surprising to ornithologists that the reversed hallux, reversed 
pubis, bony sternum, fused clavicles, and sideways-flexing hand 



are not strictly avian characters. Rather, they evolved in the 
somewhat larger, terrestrial, predaceous theropod relatives of 
birds. This suggests to Gauthier, as it has to some other evo- 
lutionary biologists, that these characters evolved in a com- 
pletely different functional context; that birds are not as distinct 
from other archosaurs as textbooks would have us believe; and 
that true "birdy-ness" is probably based on a much more re- 
stricted set of characteristics, mostly centered on flight itself 

Walter Bock, by contrast, takes an almost completely neo- 
biological view of this problem. Acknowledging the manifold 
gaps in the fossil record, and the severe limitations of fossils in 
answering questions of physiology, behavior, and metabolism. 
Bock regards the precise ancestry of birds as not critical to the 
problem of the evolution of flight. He reasons that the living 
world offers far more information about adaptations and phys- 
iology, and so suggests instead that the evolutionary biologist 
construct a "pseudophylogeny" of known living forms that might 
reasonably be functionally intermediate between successive evo- 
lutionary steps in the transition from nonflight to flight. Re- 
viewing the differences between the two major theories to ac- 
count for the origin of flight (the cursorial or "from the ground 
up" theory, and the arboreal or "from the trees down" theory). 
Bock emphasizes the theoretical difliculties with the cursorial 
model, for which no good living prototypes exist, and advocates 
the arboreal model, for which living arboreal gliders may be 
regarded as suitable prototypes. 

John Ostrom, like Gauthier, grounds his theory for the origin 
of flight (no pun intended) firmly in the phylogenetic basis for 
the origin of birds among small Mesozoic coelurosaurs. Rea- 
soning that this group of animals must have been small, agile, 
cursorial predators, and noting the complete absence in Ar- 
chacopteryx (the earliest known fossil bird) of what we would 
recognize in living animals as arboreal specializations, Ostrom 
believes it is more consistent with the available evidence to 
hypothesize that birds evolved flight from the ground up. Os- 
trom is careful to avoid saying that protobirds could not have 
gotten into trees; he merely sees no evidence for arboreality. 
This contrasts with Bock's view that whether or not protobirds 
mainly lived in trees, they must have gotten up there to launch 
themselves into the air. The diffenng consequences of these 
views are clear. Bock believes that a gliding stage was necessary; 
Ostrom does not. Bock believes that protobirds must have been 
able to climb; Ostrom does not. Ostrom favors the model of 
Caple, Balda, and Willis that shows how a running, leaping 
protobird could have developed stability in long, outstretched 
protowings, and how the resulting incremental lift and control 
could have paved the way for active, flapping flight. Bock is 
unimpressed with this model, but favors the objections of Ray- 
ner and those of Norberg that the speeds at which an animal 
would have had to run along the ground to achieve the minimum 
power speed of flight is prohibitively high to be considered 
plausible. 

The portion of the argument that hinges on characteristics 
necessary to sustain lift and motion through the air is covered 
by the preeminent biological aerodynamicist Colin Pennycuick. 
Pennycuick outlines the requirements of air travel, both aero- 
dynamic and energetic, and suggests that arboreal gliding is a 
far easier way to begin flight than any reasonable alternative. 



In his view, however, not all animals may have begun to fly in 
the same way, and he considers possible biological factors that 
may account for some of the observed differences among birds, 
bats, and pterosaurs. 

Though consensus may not have been reached here on many 
of the major problems associated with the origin of flight, there 
is no question of the importance of the problem as a paradigm 
for the understanding of major evolutionary questions. Flight 
appears to be an enormously difficult adaptation to evolve, in 
the sense that only three groups of tetrapods have ever been 
known to do it, and no nonflapping groups living today are 
obviously on their way to active flight. Yet flight is energetically 
cheaper than other modes of terrestrial transportation. If it is 
so advantageous, why is it so rare? The phylogenetic, ecological, 
and functional evidence presented in this book may provide 
some of the answers. The papers given here represent a fair 
cross-section of the approaches and opinions current in modem 
biology that have been applied to this question. The debate will 
no doubt continue; some compromises in both inference and 
method will undoubtedly surface; some problems will remain 
unsolved and rankling. This book will have done its job if it 
educates the diverse audience for which it is intended — orni- 
thologists, paleontologists, vertebrate biologists of all interests— 
in the current issues related to the origin of birds and their flight; 
and if it stimulates its readers to read and learn more about 
what we know, why we think as we do, and how our training 
influences our approaches to scientific problems. After all, the 
unsolved problems may only require a fresh look from a new 
perspective; and that, at least, is available to everyone. 

Two recent books will also be of interest to readers of this 



volume. In 1984, an international symposium on Archaeopteryx 
was convened at Eichstatt, West Germany. The proceedings of 
that symposium were collected as short papers in a volume 
entitled The Beginnings of Birds (M. K. Hecht, J. H. Ostrom, 
G. Viohl, and P. Wellnhofer, eds.), and published by the Freunde 
des JuraMuseums Eichstatt at the end of 1985. Another com- 
panion volume, for readers further interested in the aeronautical 
aspect, is \'ertebrate Flight: A bibliography to 1985. compiled by 
Jeremy M. V. Rayner (University of Bristol Press, Bristol, En- 
gland; 1985). The alphabetical bibliography is cross-indexed 
according to various topics dealing with flight and the verte- 
brates that have evolved it. These two books should serve as 
useful starting points for those interested in pursuing further the 
topics treated in the present book. 

For making the symposium and its volume possible, I would 
like to thank and acknowledge several people. Luis Baptista 
organized the American Association for the Advancement of 
Science Symposium at which these talks were presented. Daphne 
Fautin coordinated the initial editing and reviewing chores and 
provided much sensible advice and comment. Sheridan Warrick 
and Kathenne Ulnch. of the Academy's Publications Oflice, did 
a superb job of copy editing the manuscripts for clarity and 
consistency, and offered many useful suggestions. To these peo- 
ple, the reviewers, and the California Academy, the authors and 
I are most indebted. 

Kevin Padian 

Department of Paleontology 
University of California, Berkeley 
January 1986 



Saurischian Monophyly and the Origin of Birds 

Jacques Gauthier 

Museum of Zoology. Vniyersily of Michigan, 
Ann Arbor, Michigan 48109 



And if the whole hind quarters from the ilium to the toes, of a half-hatched chicken could be suddenly enlarged, ossified, and fossilized as they 
are, they would furnish us with the last step of the transition between Birds and Reptiles; for there would be nothing m their characters to prevent 
us from refemng them to the Dinosauria. 

T. H. Huxley 1870a 



Introduction 

The origin of birds has long been of interest to evolutionary 
biologists. Darwin and his colleagues were well aware of the 
problem presented by the "embarrassing gap" between birds 
and other amniotes (Desmond 1984). Indeed, because birds 
appear so different from other tetrapods, it could be argued that 
ne.xt to the question of the origin of man. that of the origin of 
birds was one of the most serious impediments to a general 
acceptance of Darwin's concept of the transmutation of species. 
The precise relationships of birds remain controversial, al- 
though the hypothesis that birds are archosaurs is now widely 
accepted. One of the goals of this report is to show that this 
controversy has much less to do with the evidence than it does 
with philosophical issues regarding interpretations of the evi- 
dence. I believe that the simplest and most interesting inter- 
pretation of the limited evidence available to T. H. Huxley and 
his contemporaries was that birds are part of Dinosauria. One 
would have hoped that work subsequent to Huxley's would have 
been directed towards determining the precise position of birds 
within dinosaurs. However, for reasons that will be discussed 
below, this was not to be the case. In fact, Huxley's views on 
bird origins were largely abandoned for much of the twentieth 
century, and the phylogenetic relationships of birds remain a 
persistent controversy. 

Among current hypotheses for the relationships of birds among 
amniotes, only three have been supported by shared apomor- 
phies and thus can be considered legitimate hypotheses in a 
phylogenetic context. Simply stated, these hypotheses are: 1) 
birds are the sister-group of mammals; 2) birds are the sister- 
group of crocodiles; and 3) birds are dinosaurs. The last two 
hypotheses agree that birds are archosaurs. The third could be 
divided into competing subhypotheses regarding the precise po- 
sition of birds within Dinosauria, but this need not concern us. 
The first two hypotheses will be briefly reviewed below, along 
with a summary of the history of the third alternative, the prin- 
cipal focus of this paper. 

Early workers suggested hypotheses, such as that birds were 
derived from early pterosaurs (Owen 1875; Seeley 1881), or 
from "lizards"' (Vogt 1879, 1880), or from"thecodonts" (Broom 
1913). These and other hypotheses of their kind are ill-advised 



' The use of quotation marks around a group name means that in this context 
that group is paraphyletic. Paraphyletic groups include an ancestor and some (but 
not all) of Its descendants (e.g., "mammals" without humans, or "lizards" without 
snakes, which are more closely related to some "lizards" than other "lizards" are). 
Monophyletic groups include an ancestor and all of its descendants, and these are 
the only kinds of groups that are considered legitimate for phylogenetic analysis. 



because they violate a basic tenet of evolutionary theory; as 
Michael Ghiselin put it (pers. comm.), "only species speciate, 
genera do not generate." Leaving aside the objection to the idea 
that higher taxa can act as ancestors in the usual sense, these 
hypotheses suffer for other reasons. 

The pterosaur-bird hypothesis never received much attention, 
and the bird-"lizard" proposition deservedly received even less 
in that it was based entirely on plesiomorphic resemblances 
(such as sharing a long tail). In light of current knowledge, many 
apomorphies shared by birds and pterosaurs are most parsi- 
moniously interpreted as convergences, which explains why the 
similarities between birds and pterosaurs are seldom either de- 
tailed or general to both taxa. However, the analysis below 
reveals some evidence that could be interpreted as supporting 
a somewhat modified version of Owen and Seeley 's hypothesis: 
namely, that pterosaurs might be most closely related to the- 
ropods as a whole and thus to the subgroup of theropods in- 
cluding birds. Martin (1983a.i 1 1 ) cited Scleromocli/us as shar- 
ing "a remarkably high percentage of the features suggested to 
relate birds to coelurosaurs." Martin evidently was unaware that 
Huene (1914a, 1948, 1956) thought that Sderomochlus might 
be related to pterosaurs. A more explicit restatement of Huene's 
hypothesis, including additional corroborating evidence, may 
be found in Gauthier (1984) and Padian (1984). 

The Thecodont-Bird Hypothesis 

One of the oldest and most popular hypotheses for the origin 
of birds has been that they evolved from "pseudosuchians" 
(hereinafter referred to as the "thecodont" ancestry hypothesis). 
"Pseudosuchians" comprise a basal group within the basal group 
of archosaurs, the "thecodonts," from which all other archo- 
saurs, including birds, are supposed to have evolved. This hy- 
pothesis has been the chief rival of the dinosaur hypothesis for 
much of the twentieth century. The "thecodont" ancestry hy- 
pothesis was proposed by workers who were impressed with two 
main objections to the dinosaur hypothesis: 1 ) "dinosaurs" were 
either morphologically too specialized or stratigraphically too 
late to have given rise to birds; 2) convergence could explain 
equally well the similarities shared by "dinosaurs" and birds. 

The first objection is not without merit, but it addresses only 
part of the issue. It emphasizes the dearth of evidence for direct 
filial relations of birds among dinosaurs, but it ignores the abun- 
dant evidence supporting common ancestry. (Just because one's 
cousins are not one's ancestors does not mean that one is not 
related to them.) 

The second objection to the dinosaur hypothesis is more sub- 



111 



ORIGIN OF BIRDS AND EVOLUTION OF FLIGHT 



tie than the first but is equally misguided. It is possible that 
birds and "dinosaurs" are convergent on one another because 
of their bipedal habits, but this is not the same as having evi- 
dence to support such a claim. Simply invoking the possibility 
of convergence is not enough (e.g., Charig 1976a, b). Both ho- 
mology and homoplasy are deductive concepts contingent upon 
accepting one of several phylogenetic hypotheses (Rieppel 1980); 
the apomorphies supporting the preferred hypothesis are con- 
sidered synapomorphies and those supporting alternatives are 
considered nonhomologous (e.g., convergent). Thus, without an 
alternative hypothesis of relationship, it is not possible to rec- 
ognize convergence. Moreover, one must accept a shared apo- 
morphy as a potential synapomorphy; to assume otherwise at 
the outset simply makes the convergence argument invulnerable 
to test. 

For example, one is able to reject the proposition that two 
taxa possessing an apomorphic, mesotarsal ankle joint shared 
a more recent common ancestor within Archosauria; one need 
only observe that a newly discovered sister-taxon of either one 
of these mesotarsal archosaurs displays the ancestral condition 
(assuming no reversals). Indeed, both the convergence and ho- 
mology propositions would agree on the significance of finding 
a member of the ingroup with the ancestral condition (i.e., con- 
vergence). However, if one simply asserts that the mesotarsal 
condition arose convergently, this proposition would be im- 
mune to new data; that is to say, no matter how many times 
newly discovered members of the ingroup were found to possess 
mesotarsal joints, one could still say that the crucial fossil (i.e., 
the unknown common ancestor) with the ancestral condition 
has yet to be found. There is only one simple and informative 
explanation for the mesotarsal joints of the birdlike archosaurs, 
and that is homology. There is, however, no limit to how com- 
plex an alternative explanation might be; taken to its logical 
extreme, it might as well be argued that all 9,000 species of 
birds acquired their mesotarsal ankles convergently. 

By the early twentieth century an apparent alternative, the 
hypothesis of "thecodont" origins, became fashionable, and ac- 
cepting this thesis required accepting the convergence argument. 
The two objections listed above may have set the stage for 
acceptance of the "thecodont" origins hypothesis, but broad 
acceptance of this idea arose with the description of one of the 
earliest and most generalized archosaurs, Euparkcna capensis. 
Broom (1913) and Heilmann (1926) then had an archosaur that 
was so ancient and primitive that it enabled them to derive 
birds directly from such a stage in archosaur evolution without 
having to pass through dinosaurs. 

If in fact birds were so related, then one would expect to find 
evidence that all dinosaurs shared a more recent common ances- 
tor with one another than any of them did with birds. Heilmann 
(1926) may have recognized this necessity, because he noted 
that the dinosaurs then known lacked clavicles. This suggested 
that birds, which have clavicles, were plesiomorphic compared 
to dinosaurs in this respect, and must have diverged from other, 
more primitive archosaurs that retained clavicles. The single 
piece of evidence supporting the "thecodont" origins hypothesis 
was refuted in 1936 when clavicles were found in nonavian 
dinosaurs (see Part IV, character 58). 

More importantly, the great weight of evidence marshalled 
by Heilmann (1926) provides clear evidence that birds are the- 
ropod dinosaurs. That is to say, even if Heilmann were correct 
in concluding that nonavian dinosaurs lacked clavicles, the weight 



of his evidence still favored the thesis that birds are dinosaurs. 
Heilmann ultimately rejected the evidence favoring the dinosaur 
hypothesis and concluded instead that birds diverged from other 
archosaurs pnor to the origin of dinosaurs. Heilmann's decision 
appears to have been predicated on a strict interpretation of 
Dollo's law of the irreversibility of evolution. Granted, one 
would not expect a hummingbird to evolve back into Archaeop- 
teryx. especially by exactly reversing the historical sequence 
leading from the latter to the former, but one stretches the point 
by concluding that if clavicles were lost then they could never 
be regained. 

The inadequacies of the "thecodont" ancestry hypothesis are 
most clearly seen in its claim that birds arose from an unknown 
member of "Thecodontia." From a phylogenetic perspective, 
"Thecodontia" and Archosauria are diagnosed by the same syn- 
apomorphies. Thus, these taxa are redundant, and when one 
says that birds evolved from "thecodonts" one is simply reit- 
erating that birds are part of Archosauria. To say that birds are 
archosaurs would be true regardless of whether or not they were 
most closely related to crocodiles, pterosaurs, omithischians, or 
any other monophyletic group within archosaurs. Thus, the 
"thecodont" ancestry hypothesis cannot be considered a legit- 
imate alternative to the mammal-bird, crocodile-bird, or di- 
nosaur hypotheses. Moreover, the "thecodont" ancestry hy- 
pothesis IS a red herring that has served to deflect interest in 
and examination of the legitimate hypotheses of bird origins. 

For example, as recently as 1980, Tarsitano and Hecht argued 
that birds were derived from "advanced thecodonts" that lacked 
dermal armor and had a birdlike ankle joint. In fact, these and 
other attributes apply to Lagosuchiis. pterosaurs, and all dino- 
saurs, including birds (see Appendix A). Tarsitano and Hecht's 
(1980:176, fig. 9) cladogram correctly depicted birds as part of 
this group, but it also made other claims that were unsupported 
by evidence. For example, their "Theropoda" and other "Saur- 
ischia" were shown to be most closely related within this group, 
with Lagosuchiis as their sister-group, and birds as the sister- 
taxon of the Lagositchus-"saur\sch\an" group. With the excep- 
tion of birds, however, they failed to diagnose these taxa and 
thus support their conclusions. A more accurate depiction of 
their results would have been a cladogram with an unresolved 
multichotomy from the level at which the characters "meso- 
tarsus" and "unarmored" arose. This would have established 
that "thecodonts" are a paraphyletic group, and that birds are 
part of the unarmored, mesotarsal group of archosaurs. One is 
still left with the possibility that birds are more closely related 
to some members of this group than they are to others. Rather 
than address this issue, however, Tarsitano and Hecht (1980: 
177) opted instead to "await better thecodont material and stud- 
ies." In so doing, they provided an example of how the use of 
paraphyletic groups as ancestors can deflect interest from evi- 
dence relevant to hypotheses of common ancestry. 

In conclusion, because the "thecodont" ancestry hypothesis 
is at best redundant and at worst a red herring. I recommend 
that It be ignored by future workers; discovering phylogenetic 
relationships among the birdlike archosaurs is difficult enough 
without the further obfuscation afforded by relying on para- 
phyletic groups as ancestors. 

The Mammal-Bird Hypothesis 

According to Desmond (1975, 1979, 1984), Owen (184 1) used 
Lamarck's three criteria of nervous, respiratory, and vascular 



GAUTHIER; SAURISCHIAN MONOPHYLI' 



organization to argue that dinosaurs, like birds and mammals, 
ascended to the physiological heights on Nature's ladder. Owen 
could hardly have been driven to such a conclusion by the scant 
remains of the three dinosaurs then known. Indeed, Desmond 
argued that by raising these huge saurians to ordinal status, 
Owen could argue that the "reptilian type" had long ago reached 
its apex, and that it had subsequently "degenerated into a sorry 
swarm of lizards" (Desmond 1984:1 19). Thus, Owen's Dino- 
sauria depended less on evidence than on his abhorrence of 
Progressivism; his ulterior motive in recognizing the taxon was 
"to add one more nail to the transmutationist coffin" (Desmond 
1979:233). 

Owen's hypothesis received little attention in the post-Dar- 
winian era, but it has been revived recently by Gardiner (1982). 
I will not here offer an extensive criticism of Gardiner's work, 
which dealt with tetrapod phylogeny in general and not just with 
the relationships of birds. Rather, I will confine myself to more 
general criticisms and let the evidence discussed below speak 
to Gardiner's (1982:227) claim that the "detailed correspon- 
dence between mammals and birds far outweighs" synapomor- 
phy schemes supporting alternative hypotheses. 

Gardiner intended to discover the relationships of "various 
living tetrapod groups ... to one another" (1982:208) through 
"consistent patterns of derived characters" (1982:228). His ac- 
tual effort was somewhat less ambitious, however, for he re- 
viewed only that evidence consistent with the conclusions of 
some pre-Darwinian comparative anatomists. Moreover, he ap- 
pears to have overlooked much of the evidence pertinent to 
tetrapod classification gathered by comparative biologists in the 
post-Darwinian era. Leaving aside these oversights, and certain 
misinterpretations of the evidence, Gardiner must be applauded 
for summarizing the apomorphies shared by mammals and birds 
that are absent in other extant amniotes; his hypothesis at least 
marshalled evidence in an explicitly phylogenetic context, unlike 
the case of the "thecodont" ancestry hypotheses. 

In view of Gardiner's sedulous pursuit of apomorphies shared 
by mammals and birds, it is curious that he uncovered only 
four shared apomorphies — the form of the heart, aortic arches, 
occipital condyle, and pattern of temporal fenestration— that 
might contradict his hypothesized close relationship between 
mammals and birds. As Gardiner's sources for this evidence 
reveal, however, his search for contrary evidence came to a 
virtual halt at the beginning of the twentieth century. Moreover, 
Gardiner claimed that he was unable to find a single unique 
feature shared by dinosaurs and birds (1982:222). In view of 
the evidence presented below, which summarizes only part of 
the relevant information already in the literature, it is difficult 
to comprehend his failure in this endeavor. Perhaps the problem 
resides in Gardiner's use of the term "unique"? But this cannot 
be the case because if one accepts the mammal-bird hypothesis, 
then endothermy is unique, but if one accepts an alternative, 
such as the crocodile-bird hypothesis, then endothermy is not 
unique. Thus, the term "unique" as used by Gardiner reflects 
a conclusion, rather than an observation, and it cannot be a 
complete explanation of his oversight. 

Perhaps Gardiner, like Owen, pursued the question of avian 
relationships with an ulterior motive. By virtually ignoring evi- 
dence determinable in both fossil and Recent taxa, and by stress- 
ing instead only part of the evidence determinable exclusively 
in extant forms, Gardiner may have revealed his intent: he was 
considerably less interested in reviewing all the evidence than 



in chastizing post-Darwinian paleontology for what he viewed 
as its excessive influence on our views of tetrapod phylogeny. 
If Gardiner had been more faithful to his avowed goals as a 
comparative biologist, he would have made a more complete 
survey of the literature upon which the evidentiary basis of the 
traditional view rests. The evidence supporting the traditional 
view that birds are archosaurian diapsids is reason enough to 
understand why comparative biologists have long held that, 
although birds and mammals are "warm in the palm of the 
hand," their immediate common ancestor among amniotes was 
not. 

The Crocodile-Bird Hypothesis 

Few have doubted that birds and crocodiles are one another's 
nearest relatives among extant amniotes. But Walker (1972, 
1974, 1977) went further by claiming that birds and crocodiles 
were most closely related even within archosaurs. Walker (pers. 
comm.) subsequently rejected this hypothesis in favor of the 
theropod dinosaur hypothesis, but the crocodile-bird hypothesis 
has received further attention from Whetstone and Martin ( 1 979, 
1981), Martin, Stewart, and Whetstone (1980), and Martin 
(1983a). Martin (1983a) listed thirty characters that had been 
used to support the crocodile-bird hypothesis, but he accepted 
Tarsitano and Hecht's (1980) argument that nearly half of them 
are plesiomorphic resemblances. Among the characters that he 
did not reject owing to obvious plesiomorphy are the following. 

1) Bipartite quadrate articulation, with apomorphic attach- 
ments anteriorly with the prootic and laterosphenoid and 
posteriorly with the prootic (in that the quadrate cotylus 
lies at the anterior base of the parocciput); 

2) Fenestra pseudorotundum carrying the perilymphatic duct; 

3) Foramen aerosum in mandible; 

4) Periotic pneumatic cavities in dorsal, central, and rostral 
positions; 

5) Two pneumatic cavities surrounding the cerebral carotid 
arteries; 

6) Pneumatic quadrate; 

7) Unserrated teeth; 

8) Tooth crowns short, bluntly conical, and with triangular 
profile; 

9) Constriction between crowns and roots of teeth; 

10) Expanded bony root covered with cementum and con- 
nected to jaw by periodontal ligaments; 

1 1) Oval or round resorption pit; 

12) Replacement tooth tilts labially and its main development 
takes place within the pulp cavity of its predecessor; 

13) Teeth implanted in open groove at least in young individ- 
uals; and 

14) Lingual walls and septa of major tooth-bearing bones formed 
by extensions of dense bone. 

With the possible exception of a foramen aerosum. at least 
some of the pneumatic sinuses in the skull, and characters 10, 
1 3, and 1 4 among those related to tooth-form and implantation, 
these characters do not appear to have been present in Archo- 
sauria ancestrally. I have observed a foramen aerosum in the 
camosaurtheropods Tyrannosaunis and Albertosaurus in a form 
essentially identical to that of extant birds (and unlike that of 
crocodilians). Other archosaurs are reported to have a croco- 
dilelike foramen aerosum. For example, Wellnhofer (1985) has 



ORIGIN OF BIRDS AND EVOLUTION OF FLIGHT 



reponed it in pterosaurs and J. Clark (pers. comm.) has observed 
this structure in the archaic archosaur Eiiparkena. Thus, the 
distribution of this character among archosaurs indicates that 
it may be the ancestral condition. However, L. Witmer (pers. 
comm.) notes that this foramen is absent in Deiuonychus. Fur- 
ther examination of other archosaurs must be undertaken to 
determine the level of synapomorphy, and subsequent history, 
of this character. 

Martin (1983a) noted that the fenestra pseudorotundum in 
birds and crocodiles can be distinguished on developmental 
novelties from analogous structures that evolved independently 
in mammals on the one hand and squamates on the other (see 
Gauthier, Estes, and de Queiroz, in prep.). Identification of a 
fenestra pseudorotundum in fossils is complicated by the ab- 
sence of soft anatomical and developmental evidence, and one 
is left only with such evidence as can be inferred from the bony 
cranium and endocasts. Unfortunately, there are few well pre- 
pared braincases preserved in a way that could provide an un- 
ambiguous conclusion regarding its presence or absence among 
archosaurs. According to Walker (1972, 1974, 1977, pers. 
comm.), fenestra pseudorotunda are absent in the extinct rela- 
tives of crocodylomorphs, the aetosaurs and parasuchians, so 
this fenestra appears to be absent in Archosauria ancestrally. 
However, Raath and Walker (pers. comm.) identified a fenestra 
pseudorotundum in the theropod Syntarsiis. I have observed a 
small fenestra in the same part of the ear region that might be 
the fenestra pseudorotundum in another member of the Syn- 
/6;/ii«'clade,Z)/7op/;twa»n« (see Ceratosauria below). This struc- 
ture also appears to be present in one carnosaur, Acrocantho- 
saunis. although it is said to be absent in the carnosaur 
rv'ra/!«05ai(n« (Whetstone and Martin 1979, 1981). However, 
L. Witmer (pers. comm.) also examined Tyraiinosaiinis and 
concluded that the fenestra pseudorotundum was present. Wit- 
mer and Martin (pers. comm.) used different criteria for rec- 
ognition of this character, so further study will be necessary to 
resolve this issue. Fenestra pseudorotunda are also reported in 
troodontid, omithomimid. and caenagnathid theropods (e.g., 
Barsbold 1983; Currie 1 986a. b), all of which appear to be closer 
to birds than are either Syntarsus-Dilophosaums or camosaurs 
(see Coelurosauria below). Whetstone and Martin (1981) ques- 
tioned these identifications, and not without reason, for it is 
difficult to distinguish this fenestra from others connected to the 
more (e.g., caenagnathid) or less (e.g., omithomimid) extensive 
periotic pneumatic cavities in the crania of these theropods. 
Whetstone and Martin (1979, 1981) were unable to identify a 
fenestra pseudorotundum in an ankylosaur omithischian or in 
hadrosaurian omithopod omithischians. However, Sues (1980) 
and Gallon (1983) claimed that it was present at least in or- 
nithopods ancestrally. Thus, there appears to be some question 
regarding the level(s) at which this synapomorphy arose within 
Archosauria. Finally, characters 1, 2, 4, 5, 9, and 13 do not 
appear to support an exclusive relationship between crocodiles 
and birds within Archosauria, because Currie (1986a, b) has 
recorded their presence in the theropod Troodon (=Stenony- 
chosawus). 

Martin (1983a) noted that no one has argued that the ancestry 
of birds lies within crocodiles, only that they both share an 
"unknown common ancestor." Martin (1 983a.- 1 1 1) argued that 
the crocodylomorph Sphenosuchus is not "as similar to birds 
as is the Crocodilia"; the common ancestor of birds and croc- 



odiles, as so proposed, must lie within Crocodylomorpha. Un- 
fortunately, this relationship has not been made explicit; birds 
have yet to be placed within a cladogram depicting their precise 
relationship among crocodylomorphs. Such an analysis must 
eventually be undertaken by the proponents of the crocodile- 
bird hypothesis in order to make their hypothesis more ame- 
nable to test. For example, in a preliminary phylogenetic anal- 
ysis of crocodiles and their extinct relatives, Crocodylomorpha 
was diagnosed by 1 3 synapomorphies (Gauthier 1 984). Of these, 
only 7 are present in birds, indicating that if birds are croc- 
odylomorphs, the former diverged from within the latter after 
the acquisition of the first seven shared apomorphies but before 
the remaining 6 crocodylomorph characters appeared. More- 
over, the 14 additional characters diagnostic of true crocodiles 
within Crocodylomorpha must have evolved later still. This 
raises questions regarding Martin's interpretations of the level 
of synapomorphy of several characters, among them being the 
characters in tooth form and implantation cited above. That is 
to say, the tooth form unique to crocodiles appeared within 
crocodylomorphs only after the evolution of an anterodorsally 
inclined quadratojugal that extends to the skull roof, a partial 
secondary palate formed by the maxillae, loss of the ventral 
process of the squamosal, loss of clavicles, ventromedial elon- 
gation of the coracoid, and development of a characteristic wrist 
joint involving an elongate and columnar radiale and ulnare. 
However, some crocodylomorphs, such as Terrestrisuchus (Crush 
1984), possess all six of the characters listed above in addition 
to those shared by birds and other crocodylomorphs, indicating 
that it is closer to true crocodiles than are birds. The problem 
emerges in the presence of sharply pointed, serrated teeth in 
Terrestrisuchus that appear to be implanted and replaced as in 
Archosauria ancestrally. If, as these data suggest, Terrestrisuchus 
and true crocodiles are closer to one another than either is to 
birds, then the dental apomorphies shared by birds and part of 
Crocodylomorpha must have been acquired convergently. This 
class of problems faces several of the supposed bird-crocodile 
characters listed above. In order to recognize and come to grips 
with such cases of character discordance, however, one requires 
a more precisely stated hypothesis than that birds and crocodiles 
share "some unknown common ancestor" (Martin 1983a. 1 1 1), 
because this would be true in any case under the traditional 
view that birds are archosaurs. 

The Dinosaur Hypothesis 

Huxley's dinosaur hypothesis found its roots in Haeckel ( 1 866), 
who considered birds to be basically "reptilian," and in earlier 
work of his own (Huxley 1864, 1867) in which he concluded 
that birds were derivatives of "sauropsid reptiles." Huxley was 
joined in supporting a "dinosaurian" origin of birds by other 
comparative anatomists including Cope (1867), Schmidt ( 1874), 
Marsh (1 877), Gegenbaur( 1878), Williston (1879), W. K. Par- 
ker(1864), T. J. Parker (1882), Baur(1883, 1884, 1885, 1886), 
and Darwin (1872). Indeed, Huxley's proposed "dinosaurian 
affinities" of birds gained broad acceptance during the latter 
quarter of the nineteenth century, even among the ranks of those 
who where "at best indifferent to Darwin" (Desmond 1984: 
132). 

In Huxley's 1869 address before the Geological Society, he 
described the "ornithic peculiarities" of Dinosauria as opening 



GAUTHIER: SAURISCHIAN MONOPHYLY 



up "a very interesting field of inquiry" that inspired him to 
devote "all my disposable leisure during the winter of 1867-8" 
to discovering characters shared by "dinosaurs" and birds that 
are not also shared by "lizards" and crocodiles. During this time 
Huxley also set out to examme critically "the material in the 
British Museum in order to ascertain how far the peculiarities 
of Megalosaurus were common to the Dinosauria in general." 
Huxley (1868, 1870a, ft) cited 35 characters as "evidence of the 
affinity between dinosaurian reptiles and birds." Of these the 
following 1 7 characters survive critical examination in light of 
our current knowledge. 

1) The skeleton is hollow and lightly constructed. 

2) The cervical vertebrae are elongate. 

3) There are more than two sacral vertebrae. 

4) The scapula is elongate and narrow. 

5) The coracoid is short and rounded. 

6) The ilium is prolonged anteroposteriorly. 

7) The acetabulum is roofed above by a supracetabular but- 
tress of the ilium. 

8) The bony contribution of the ilium to the acetabulum is 
more or less replaced by membrane. 

9) The ischium and pubis are much elongated. 

10) The femur has a strong anterior trochanter. 

1 1) The femur has a crest on the ventral face of the outer con- 
dyle that passes between the tibia and the fibula. 

12) The proximal end of the tibia is produced anteriorly into 
a strong crest, which is bent outwardly, or towards the 
fibular side. 

13) There is a crest on the lateral side of the proximal end of 
the tibia for attachment of the fibula. 

14) The tibia has a fossa distally for the reception of the as- 
cending process of the astragalus. 

15) The fibula is gracile compared to the tibia, and its distal 
end IS much smaller than the proximal. 

16) The astragalus is compressed, its articulation with the tibia 
is concave proximally and it has a convex, pulleylike distal 
surface, and the disparity in size between the tibia and fibula 
is also reflected in the astragalus being much larger than 
the calcaneum. 

17) An "ascending process" (the intermedium) more or less 
tightly connects the astragalus and tibia. 

Only some of these characters can now be considered to have 
arisen in the immediate common ancestor of Dinosauria (see 
Appendix A). Nevertheless, all of them appeared within the 
subgroup of archosaurs containing birds and thus they speak 
against the crocodile-bird hypothesis. At first glance, the ex- 
planatory powers of the crocodile-bird versus dinosaur hypoth- 
eses do not appear to di ffer greatly in that the former is supported 
by 14 shared apomorphies and the latter by 17. One must bear 
in mind, however, that the dinosaur hypothesis was formulated 
over a century ago with a fraction of the evidence currently 
available (see Appendix A). Although the crocodile-bird hy- 
pothesis is based on current knowledge, there would have been 
no reason to accept it over the dinosaur hypothesis as it stood 
in 1870, and there is even less reason to do so now. Huxley 
(1870/1) mistakenly claimed that the absence of clavicles was 
diagnostic of Dinosauria, but this did not deter him from hy- 
pothesizing a bird-"dinosaur" affinity. I doubt that he consid- 
ered the matter in this way, but he was correct in his judgement 



to the extent that it would have been simpler to accept the 
reappearance of clavicles in birds than to accept convergence 
as an explanation for each of the seventeen characters listed 
above. 

In the discussion following Huxley's presentation before the 
Geological Society in 1 869, Seeley remarked that he "thought 
it possible that the peculiar structure of the hinder limbs of the 
Dinosauria was due to the functions they performed rather than 
to any actual affinity with birds" (Huxley 1870a:31). With this 
simple declaration arose the issue that was to become the bane 
of Huxley's hypothesis. Seeley suggested convergence as an al- 
ternative explanation for the apomorphies shared by "dino- 
saurs" and birds without proposing the requisite alternative 
hypothesis of relationship. 

The assertion of convergence was to be heard time and again 
in the ensuing years (e.g., Mudge 1879; Dollo 1882, 1883; Dames 
1884; Parker 1887; Furbringer 1888; Osbom 1900). What made 
matters worse was the rise of the "thecodont" ancestry hypoth- 
esis in the early part of the twentieth century (e.g.. Broom 1913; 
Heilmann 1926); this hypothesis appeared to dignify the oth- 
erwise bald assertion of convergence. The error was compound- 
ed even further when the convergence argument was applied 
not only to bird origins but to the question of dinosaur mono- 
phyly as well. Pandora's box had been opened; if an erect, bi- 
pedal posture and gait arose convergently in dinosaurs and birds, 
then what was to forbid multiple evolutions of such adaptively 
significant characters? The demise of the dinosaur-bird hypoth- 
esis went hand in hand with the demise of dinosaur monophyly; 
if one accepted the nonparsimonious reasoning behind the hy- 
pothesis that birds were "derived independently" from "thec- 
odonts," then why not accept the even less parsimonious hy- 
pothesis that each of the remaining dinosaurian groups were 
"derived independently" from "thecodonts" as well? I see noth- 
ing that would forbid the multiple evolution of "dinosaurian" 
characters, but what evidence indicated that this did in fact take 
place? Although a few researchers advocated the dinosaur hy- 
pothesis (e.g., Boas 1930), the "unknown ancestor," coupled 
with the convergence argument, enthralled most systematists 
(e.g., Simpson 1946; de Beer 1954; Romer 1966). 

There was renewed interest in the origin of birds in the 1 970's, 
beginning with the publications of Galton (1970), Walker (1972), 
and Ostrom (1973). Galton ( 1 970) stressed one of the characters 
initially employed by Huxley (1870a, b)—a reversed pubis— to 
suggest that birds were "derived" from omithischian dinosaurs. 
Upon introduction to Ostrom's ( 1 973) evidence, Galton rejected 
the omithischian hypothesis (e.g., Bakker and Galton 1974). 
Martin ( 1 983a) suggested that the presence of a predentary bone 
in Hesperomithes and its inferred presence in Ichthyornis may 
be further evidence of an omithischian-bird relationship. The 
absence of a predentary bone in Archaeopteryx and all other 
birds speaks against Martin's conclusion; even if omithischians 
were the sister-group of birds, it would still be simpler to accept 
a predentary bone as diagnostic of a taxon including only Ichthy- 
ornis and Hesperomithes. 

Just as Eiiparkeria spurred interest in the "thecodont" an- 
cestry hypothesis, Ostrom's finds of a new and strikingly birdlike 
theropod, Dei nonychus aiUirrhopus {OstTom 1969a. b. 1974/), 
1916b), revitalized the dinosaur hypothesis. Ostrom's (1973, 
1974a, 1975a, b. 1976a) hypothesized "coelurosaurian" ances- 
try of birds could be viewed as an extension of Huxley's pro- 



ORIGIN OF BIRDS AND EVOLUTION OF FLIGHT 



posal, since the theropod Megalosaurus figured prominently in 
Huxley's arguments. The two hypotheses differed, however, in 
that Huxley looked at "dinosaurs" in general as "intermediate" 
between "reptiles" and birds, while Ostrom sought the ancestry 
of birds among "coelurosaur theropods" alone and skirted the 
question of dinosaur monophyly. 

Early workers found favorable comparisons between birds 
and "coelurosaurs," and these comparisons were ably reviewed 
by Heilmann (1926). Although Heilmann has been considered 
the champion of the "thecodont" hypothesis, his closing com- 
ment (1926:185) upon finishing his comparisons between birds 
and "coelurosaurs" was: "We have therefore reasons to hope 
that in a group of reptiles closely akin to the Coelurosaurs we 
shall be able to find an animal wholly without the shortcomings 
here indicated for a bird ancestor." 

After Heilmann (1926), a few authors advocated a special 
bird-"coelurosaur" connection (e.g., Lowe 1935, 1944a. b: 
Holmgren 1955). As Ostrom ( 1 976a) noted, however, their ideas 
received little attention because they saddled themselves with 
burdens — most notably avian polyphyly — that inspired others 
to reject their ideas out of hand (e.g., Simpson 1946; de Beer 
1954). Ostrom's resurrection of the "coelurosaur" hypothesis 
met with wider acceptance (e.g., Bakkerand Galton 1974; Thul- 
bom 1975, 1984; Thulbom and Hamley 1982), although his 
hypothesis was not without critics (e.g.. Walker 1977; Tarsitano 
and Hecht 1980; Martin 1983a). Many of the criticisms were 
centered on the interpretation of certain anatomical details in 
imperfectly preserved fossils. Perhaps more troubling was the 
continued reliance on the part of Ostrom's critics on the "the- 
codont" hypothesis and its henchman, convergence. 

Ostrom considered birds to have "evolved from coeluro- 
saurs." However, in a phylogenetic perspective Ostrom's use of 
"coelurosaur" is no more informative than the name Therop- 
oda, because both names are diagnosible by the same synapo- 
morphies (see Introduction to the Basic Taxa, below). Thus, the 
"coelurosaur" hypothesis was no more precisely stated than was 
the crocodile-bird hypothesis, in that it merely claimed that 
birds and "theropods" shared an unknown common ancestor. 
In failing to let his hypothesis accurately reflect the structure of 
his evidence, Ostrom opened himself to a variety of criticisms, 
most of which were only misinterpretations invited by the obfus- 
cation of paraphyly. It is inappropriate to consider each of these 
criticisms at this time, and the reader is referred to the character 
discussions below for examples of the problems stemming from 
treating paraphyletic and monophyletic taxa as if they possessed 
the same properties (i.e., a unique history involving origin, di- 
versification, extinction, etc.). 

Another result of Ostrom's vague conclusions regarding the 
relationships of birds to other theropods was that some workers 
mistakenly concluded that birds and theropods might be sister- 
groups (e.g., Tarsitano and Hecht 1980; Thulborn and Hamley 
1982). The imprecision in Ostrom's proposition was rectified 
by Padian (1982), who extracted the relevant evidence from 
Ostrom's works and arrayed it in an explicitly phylogenetic 
context. Padian's provisional reanalysis demonstrated that birds 
are more closely related to some "coelurosaurs" than they are 
to others; the conclusion that birds and theropods are sister- 
groups could not be extracted from Ostrom's evidence. On the 
contrary, birds are deeply imbedded within Theropoda, just as 



humans are deeply imbedded within the phylogenetic nexus of 
Mammalia. 

Bakker and Galton (1974) integrated the hypotheses of Os- 
trom and Huxley and resurrected the concept of dinosaur mono- 
phyly. Their method of analysis was somewhat more rigorous 
than the Simpsonian "evolutionary systematics" of Ostrom, 
although they were also hampered by such paraphyletic taxa as 
"thecodonts" and "prosauropods." The essence of Bakker and 
Galton's (1974) argument was as follows. 

1) Dinosaurs (including birds) shared apomorphies not also 
shared by "thecodonts" and were therefore monophyletic. 

2) "Prosauropods" were a primitive grade of dinosaurs that 
bridged the gap between omithischian and "saurischian" 
dinosaurs. 

3) All dinosaurs, like modem birds, were active endotherms; 
the behavioral and physiological apomorphies shared by di- 
nosaurs are important; Dinosauria should therefore be ac- 
corded Class status. 

Bakker and Galton's paper inspired considerable controversy 
(e.g., Thulborn 1975; Chang 1976/>). The most telling criticisms 
were directed to the second and third points. To be sure, the 
link between Bakker and Galton's evidence and the conclusion 
of endothermy was a bit tenuous. The question of categorical 
rank of Dinosauria is trivial; current methods of establishing 
taxonomic rank rely entirely upon the authority of a system- 
alist's subjective notion of what constitutes an ideal Class, Or- 
der, etc., and such Platonic and typological notions are anach- 
ronistic at best. Charig's ( 1 976/i) criticisms of Bakker and Galton's 
second point were for the most part trenchant and insightful. 
However, although some of Thulborn's (1975) and Charig's 
(1976/') criticisms of the evidence supporting dinosaur mono- 
phyly were factually accurate, for the most part they did not 
bear on the question of dinosaur monophyly. 

Most criticisms of the resurrected Dinosauna hypothesis fell 
into one of three classes: 1 ) saurischians and omithischians are 
too different to be monophyletic, 2) some of the alleged dinosaur 
characters were in fact present in some "thecodonts," and 3) 
the rest of the characters are functionally related and they could 
have been acquired convergently. The first class of criticisms is 
beside the point. One may be different from one's siblings, yet 
still share the same parents; the differences between bats and 
whales do not preclude their being mammals any more than the 
differences between the postdentary bones of Ophiacodon and 
Homo indicate nonhomology. Criticisms in the second class 
only indicate that some "thecodonts" are closer to dinosaurs 
(including birds) than are others; thus, "Thecodontia" is para- 
phyletic. Such evidence may not support dinosaur monophyly, 
but it does not speak against it either. The third class of criticisms 
leveled at Bakker and Galton's evidence for dinosaur mono- 
phyly is exemplified by a quote from Chang (1976/).'79): "Most 
of these dinosaurian character-states were obviously adapta- 
tions to the fully improved ('fully erect') posture and gait of the 
dinosaurs . . . which could easily have evolved several times 
over, in slightly different ways, in response to similar functional 
requirements." 

We have now come full circle; the exchanges between Huxley 
and Seeley. and between Bakker and Galton on the one hand 
and Chang and Thulbom on the other, demonstrate that the 



GAUTHIER: SAURISCHIAN MONOPHYLY 



dinosaur controversy has not altered on this issue in over a 
century. This controversy has nothing to do with evidence, al- 
though it must be admitted that until Hennig 1966, it was not 
widely understood that ancestry, rather than overall similarity 
or dissimilarity, must be the basis of phylogenetic classification. 
Consequently, there is little to be gained from taking the position 
that the problems will be solved by finding more fossils. There 
are, after all, many more fossils now available than there were 
in Huxley's time, and they still have not forestalled the old 
objections. The implications are clear: if our understanding of 
bird origins is to progress, we ought to rid ourselves of typo- 
logical thought, namely, paraphyletic "Thecodontia"; reserve 
hypotheses of convergence for cases of character discordance, 
without which there is nothing for the concept of convergence 
to explain; and remember that although the world has no ob- 
ligation either to be simple or informative, our hypotheses had 
better be both (Beatty and Fink 1979). By following these pre- 
cepts, issues in archosaur phylogeny can be brought into sharper 
focus. 

Knowledge of theropod anatomy has increased vastly since 
Buckland described Megalosaunis in 1824. Unfortunately, this 
knowledge has not been translated into a deeper understanding 
of theropod phylogenetic relationships. This circumstance re- 
flects, in part, that Theropoda has yet to be diagnosed on the 
basis of synapomorphies. Indeed, most workers have been con- 
tent to define "theropods" as "primitive dinosaurs" or "car- 
nivorous saurischians"; which is to say that theropods are those 
saurischians that are not sauropodomorphs, and saurischians 
are those dinosaurs that are not omithischians. In order to bring 
these taxa into the phylogenetic system, and thereby address the 
question of the phylogenetic relationships of birds, it will first 
be necessary to determine which, if any, phylogenetic entities 
within Dinosauria might be parts of Saurischia and Theropoda. 

Materials and Methods 

For the most part, the specimens examined are listed in the 
Introduction to the Basic Taxa. However, the materials consti- 
tuting the core of the analysis were Segisaiinis. Dilophosaurus, 
casts of the skull ofCeralosawus. and several undescribed spec- 
imens in the collections of the University of California Museum 
of Paleontology (UCMP); Coelophysis in the collections of the 
Museum of Northern Arizona (MNA), UCMP, and American 
Museum of Natural History (AMNH); Procompsognathus in 
the collections of the Staatliches Museum fiir Naturkunde, Stutt- 
gart (SMNS); Compsognathus in the collections of the Bayer- 
ische Staatssammlung fiir Palaontologie und historische Geo- 
logic, Munich (BSP); and the comparative osteological collections 
of extant birds housed in the UCMP, the Museum of Vertebrate 
Zoology, University of California, Berkeley (MVZ), the Cali- 
fornia Academy of Sciences (CAS), and the University of Mich- 
igan Museum of Zoology (UMMZ). These data were supple- 
mented by photographs and notes on virtually every taxon 
referred to nonavian theropods compiled by Samuel Welles and 
Robert Long. Information on character ontogenies was derived 
from the literature and from cleared and double-stained series 
oi Alligator. Gallus. and Podiceps occidentalis (UMMZ), in ad- 
dition to late embryos and juveniles of a few passerines (CAS 
and pers. coll.), and the hindlimbs of three tinamou embryos 



(MVZ). These data were supplemented by that derived from 
skeletons of a hatchling Plerocnemia (MVZ) and a juvenile Ap- 
teryx (UMMZ). 

For the purposes of this work, ontogenies will be divided into 
four stages: embryos, juveniles, subadults, and fully adult/ma- 
ture individuals. Embryos refer to prehatching individuals; ju- 
veniles include stages from hatchlings to nearly full-grown spec- 
imens (i.e., subadults); subadults are those individuals that are 
near to maximum size as is indicated by some, but not all, of 
the developmental events marking the cessation of growth (i.e., 
senility). As used here, the term "fully adult" makes no reference 
to sexual maturity, which may or may not be coincident with 
this stage in skeletal ontogeny. Individuals that have reached 
the terminal stage of ontogeny are referred to variously as fully 
adult, fully mature, or as having attained maximum adult size. 
Subadults may also be near or at maximum adult size, but they 
do not display the full suite of developmental events in the 
skeleton that mark the cessation of growth. The cessation of 
growth in theropods may be recognized by the following events 
in the skeleton: fusion between the axial intercentrum and at- 
lantal centrum, and fusion of this compound structure to the 
axial centrum; fusion of neural arches to centra; fusion of the 
vertebral components of the sacrum; full ossification of distal 
tarsal II; and, at least in nonavian theropods aside from ratites, 
fusion of the scapula and coracoid. Some of these events may 
precede others; however, any specimen in which all of them are 
present is here considered to have attained maximum adult size. 
By this definition, there are few nonavian theropod fossils that 
have achieved maximum adult size (e.g.. the type specimens of 
Syntarsiis rhodesiensis and Cemtosawus nasiconus). There are 
also few specimens that could be considered juveniles, and even 
these few represent approximately half-grown individuals (e.g., 
the type specimen of Compsognathus longipes and. contrary to 
Howgate's 1984 view, the Eichstatt specimen oi Archaeopteryx 
lithographica). 

The methods employed here are essentially those of Gauthier 
et al. (in prep.). Following the procedure of Maddison et al. 
(1984). at least two outgroups were employed to identify 84 
apomorphic characters distributed among 2 or more of the 17 
basic taxa (18 including outgroup) that were the subjects of this 
analysis (see Introduction to the Basic Taxa, Fig. 7, and Ap- 
pendix B). These data were then analyzed with Swofford's Phy- 
logenetic Analysis Using Parsimony (PAUP) program installed 
in the University of Michigan's Terminal System. The analysis 
was two-part; the first run included only the comparatively well 
known taxa whose interrelationships were the principal foci of 
this analysis (viz., Omithischia. Sauropodomorpha, Ceratosau- 
ria, Camosauria, Omithomimidae, Deinonychosauria, and 
birds), and the second run included all taxa, including those for 
which we have relatively little information owing to incomplete 
preservation. The first run yielded a single cladogram depicting 
the most parsimonious interpretation of the phylogeny of the 
seven well-known dinosaur taxa (Fig. 8). This cladogram re- 
quires 94 evolutionary events to account for the distribution of 
84 apomorphies among the seven taxa, thus yielding a consis- 
tency index of 89%; that is to say, the single tree so obtained 
represents a highly corroborated hypothesis of relationship. 
However, the second run, which included both the better known 
and less well known theropods, resulted in numerous, equally 



ORIGIN OF BIRDS AND EVOLUTION OF FLIGHT 



parsimonious trees. Nevertheless, the sister-group relationship 
among the seven well-known taxa were consistent across all 
possible trees. The multiple trees obtained in the second run 
resulted from the missing data in the 10 less well known taxa. 
For example, let us say that Coelurosauria includes only three 
taxa: deinonychosaurs, birds, and Hulsanpes perlei. Only three 
of the 84 characters under review were determinable for Hul- 
sanpes, and although these data indicate that it is a coelurosaur, 
they are moot on the point of its precise affinities within this 
taxon. The PAUP analysis is so constructed that it considers all 
possible positions that Hulsanpes could have among coeluro- 
saurs (viz., it could be the sister-group of birds, deinonycho- 
saurs, or a deinonychosaur-bird group) and thus yields three 
equally parsimonious cladograms for these three taxa. Of course, 
none of these cladograms is actually supported by any evidence 
observable in Hulsanpes. Thus, when one considers the added 
possibilities allowed by the very incomplete remains of the 10 
less well known taxa, the source of the numerous possible trees 
becomes apparent. However, two critical points must be borne 
in mind: first, the relationships among the seven taxa for which 
there is more complete information remained unchanged, and 
second, the overwhelming majority of possible trees were in fact 
uninformative. Consequently, they were collapsed into a single 
consensus tree incorporating multichotomies stemming from 
the levels supported by observable characters (Fig. 9). The con- 
sensus tree required 99 evolutionary events to account for the 
distnbution of 84 apomorphies among the 17 basic taxa, thus 
yielding a consistency index of 85%. 

Following Gauthier et al. (in prep.), the classification used in 
this work was constructed according to the following five con- 
ventions. 

1) Only monophyletic taxa including an ancestor and all of 
its descendants are recognized, and in no case will a demon- 
strably paraphyletic taxon be considered in this analysis. An- 
cestry, rather than overall similarity, must be the basis for a 
phylogenetic system. The single exception to this convention is 
the metataxon. 

2) A new category, the metataxon, is employed for taxa for 
which there is no positive evidence for or against recency of 
common ancestry. Following the suggestion of M. Donoghue, 
metataxa are provisionally allowed in the classification and their 
uncertain status is denoted by an asterisk following the name. 
For example, as will be argued below, the monophyly or para- 
phyly of the five fossil skeletons and a single feather impression 
referred to the earliest bird. Archaeopteryx litliographica*. has 
yet to be firmly established, and it is therefore accorded meta- 
taxon status. 

3) Certain widely used names are standardized by restricting 
them to taxa whose monophyly among extant amniotes is firmly 
established. Accordingly, Archosauria is standardized by lim- 
iting this taxon to all the descendants of the most recent common 
ancestor of extant birds and crocodiles. And Aves is likewise 
restricted to all the descendants of the most recent common 
ancestor of Ratitae, Tinami, and Neognathae. 

4) Although the spelling of current taxonomic names is re- 
tained, no formal categoncal ranks are recognized and hierar- 
chical relationships within taxa are expressed instead by branch- 
ing diagrams. Categorical ranks such as Class, Order, Family, 
and Genus, will not be recognized in this work. 

5) Except to preserve binomials, no redundant names will be 



recognized. Thus, although Archaeopteryx lithographica* is re- 
tained, redundant taxa conveying nothing further about phy- 
logenetic relationships, such as Archaeopterygidae, Archaeop- 
terygiformes, or Saururae will be ignored. 

Introduction to the Basic Taxa 

This section defines and diagnoses the theropod taxa that are 
the subjects of the present analysis (For diagnoses and defini- 
tions of the other basic taxa, Sauropodomorpha and Omithi- 
schia, and the outgroup taxa, see Appendix A). These diagnoses 
are not definitive; the basic taxa are assumed to be monophy- 
letic, and the characters are listed merely to show that there is 
at least some basis for the assumption of monophyly in each 
case. 

Several workers have described the conventional groupings 
of "Camosauria" for large theropods and "Coelurosauria" for 
small theropods as inadequate in view of the observed variation 
among Theropoda (e.g., Colbert and Russell 1969; Ostrom 
1969ft). Accordingly, modem classifications emphasize less in- 
clusive taxa, typically ranked as "families." and these units will, 
for the most part, serve as the basic taxa of this analysis. In a 
phylogenetic context, "Coelurosauria" and "Theropoda" are 
redundant in that they have traditionally been diagnosed by the 
same synapomorphies. Huene (\9\Ab) originally defined "Coe- 
lurosauria" on the basis of plesiomorphic resemblances, such 
as their small size and long necks. In other words, they are 
theropods that are not camosaurs. However, because Huene 
based the concept on "coelurids," some of which have synapo- 
morphies of a particular subgroup of Theropoda that includes 
birds, Coelurosauria will be retained in a modified form (see 
below). 

A principal goal of this work is an examination of the phy- 
logenetic relationships within Theropoda, with particular atten- 
tion to the relationships among the better known theropods here 
included in Ceratosauria (n. comb.), Camosauria (n. comb.), 
Ornithomimidae (n. comb.), Deinonychosauria, and birds. For 
the sake of completeness, the distribution of the characters dis- 
cussed will also be noted in less well known taxa such as Pro- 
compsognathus iriasicus*. Liliensternus liliensterni*, Ornitho- 
lestes hermanni*. Coelurus fragilis*. Compsognathus longipes. 
Microvenator celer*, Saurormtholestes langstoni*, Hulsanpes 
perlei. Elmisauridae*, and Caenagnathidae. As the use of the 
asterisk indicates, several of these taxa have not been adequately 
diagnosed; they may indeed be different from other theropods, 
but until they are diagnosed on the basis of characters relevant 
to the question of monophyly (i.e., synapomorphies), their status 
as phylogenetic entities must remain suspect. The position of 
these generally poorly known taxa are not so critical to the goals 
of this analysis, and the reader is referred to the literature for 
more information (Marsh 188 la,- Osborn 1903, 1917; Ostrom 
1970, 1978, 1981; Sues 1978; Osmolska 1981, 1982; Barsbold 
1983; Welles 1984). 

The more inclusive basic taxa may be newly recognized, or 
they may differ in diagnosis and content from concepts em- 
ployed by other researchers. Some of the synapomorphies I 
consider diagnostic of the basic taxa could only have been rec- 
ognized as such after the completion of the analysis; they are 
added now for the sake of completeness and in no case would 
the monophyly of any of the basic taxa depend on such deter- 
minations. 



GAUTHIER: SAURISCHIAN MONOPH'lLV 



Elmisauridae* Osmolska, 1981 

Temporal Range. — late Cretaceous. 

Included J fAA. — Chiroslenoles pregmcilis*. Matrophalangia canadensis*, and 
Elmisaunts rants*. 

Diagnosis. — Based on personal observation of Macropha- 
langia* and Cliirostenotes*, and on published descriptions of 
these taxa in Gilmore (1924), Sternberg (1932), and Osmolska 
(1981). These taxa are represented by scant and often noncom- 
plementary remains. Indeed, none of the referred taxa have been 
adequately diagnosed, and they are so poorly known that Os- 
molska (1981) suggested that they might be synonymous. Ac- 
cording to Cume (pers. comm.), however, Elmisaunts* and 
Cliirostenotes* are sympatric in Alberta in the late Cretaceous. 
Currie and Russell are in the process of describing a partial 
skeleton, including hands and feet, of Chirostenotes*. Prelimi- 
nary results indicate that Chirostenotes* and Macrophalangia* 
might be based on the same species. This conclusion is tentative 
because there appear to be two "morphs," and it is not yet clear 
if these morphs result from sexual or taxonomic differences. To 
further complicate matters, each of the apomorphies shared by 
elmisaurids* is either matched in some other theropods, or it 
could be considered part of a transformation series that is taken 
to extreme in some group of theropods (e.g., proportions of 
manal digit I approach those of omithomimids). More evidence 
will be necessary to address these issues, but for the present 
Osmolska's Elmisauridae* will be accepted as a metataxon. Fol- 
lowing is a list of apomorphies shared by elmisaurids*; as sug- 
gested above, these apomorphies may or may not prove to be 
synapomorphies. 

Metacarpal I elongate and slender, relatively elongate first 
and second phalanges of manal digit III; metatarsus elongate 
and narrow; metatarsal III pinched between metatarsals II and 
IV, the latter two contacting one another proximally in front of 
III (similar modifications of the hands and tarsus are present in 
omithomimids, troodontids, and omithurine birds). 

Caenagnathidae Sternberg. 1 940 

Temporal Range. — late Cretaceous. 

Included J^XA. — Caenagnathus collinsi. C. sternbergi. and Ovirapioi philo- 
ceratops. 

Diagnosis.— No one doubts the monophyly of these pecu- 
liarly specialized theropods (Osbom 1924^; Osmolska 1976; 
Barsbold 1983). Caenagnathids were once thought to be related 
to omithomimids because both share edentulous, beaklike snouts, 
but more recent work suggests otherwise (Barsbold 1983, and 
see below). Caenagnathids have highly modified skulls, and there 
is very little information regarding their postcranial skeletons. 
Several new specimens have been discovered, but they have yet 
to be completely described and illustrated (Osmolska 1976; 
Barsbold 1983). Barsbold is currently engaged in a revision of 
this taxon based on new material including as many as three 
species of Oviraptor. and an adequate diagnosis of this taxon 
must await his findings. 

Currie (pers. comm.) has informed me that Caenagnathus 
(known only from cranial material) and Chirostenotes* (known 
only from postcranial material) might represent the same species. 
Moreover, Wilson and Currie (pers. comm.) have suggested that 
Microvenator* might be a caenagnathid. These hypotheses are 
only tentative, but they are included because Currie and Wil- 



son's observations indicate that Microvenator*, elmisaurids*, 
and caenagnathids might be monophyletic; such possibilities 
should always be bome in mind when dealing with metataxa. 

Ceratosauria Marsh, 1884ft (n. comb.) 

Temporal Range.— late Tnassic lo late Jurassic. 

Included JAXA. — Ceratosaurm nasicorms. Synlarsus rhodesiensis. Coelophysis 
bauri. Scgisaurus halh*. Sarcosaunis woodi*. Ddopliosaunis welherelli (including 
UCMP 37302, 37303, and 77270), and some undescnbed forms represented by 
UCMP 129618 (referred lo Coelophysis by Padian, m press). UCMP 128659, and 
MNA V. 2623 (referred to Syntarsus by T. Rowe, pers. comm.). 

Diagnosis. — The initial basis for recognition of the mono- 
phyly of this taxon stemmed from Welles's (1984) observation 
that one specimen referred to Dilophosaiirus (UCMP 77270) 
possessed a uniquely modified trochanteric shelf (=modified 
anterior trochanter: see photograph of Sarcosaunis woodi* in 
Charig 1976ft). T. Rowe later observed this apomorphy in Segi- 
saurus*, and we have since observed this and other shared apo- 
morphies in all taxa here included in Ceratosauria. 

The presence of the trochanteric shelf in only some ceratosaur 
specimens is perplexing. However. Colbert, Rowe, and Raath 
(pers. comm.) have separately observed the presence of two 
femoral types among the large series of Coelophysis and Syn- 
tarsus. a robust form in which the trochanteric shelf is developed 
in the form characteristic of ceratosaurs, and a gracile form in 
which the trochanteric shelf is less modified and more like that 
seen in dinosaurs ancestrally. Dimorphism in femoral form, 
along with other differences in proportions, have been attnbuted 
to sexual dimorphism. 

Although it has appeared elsewhere in theropods, another 
synapomorphy of Ceratosauria is the fusion between distal tar- 
sals 2 and 3 and their respective metatarsals (T. Rowe, pers. 
comm.; Raath 1969). Rowe (pers. comm.) has discovered ad- 
ditional synapomorphies of Ceratosauria, including the shape 
and prominence of the supracetabular shelf, a fibular groove on 
the proximal end of the lateral side of the fibula (e.g., Gilmore 
1920, fig. 65C), and a prominent groove on the ventrolateral 
side of the fibular condyle of the femur. Rowe (pers. comm.) 
also noted that, with the possible exception of Segisaurus*, all 
ceratosaurs have a narrower pubis than is seen in other thero- 
pods aside from birds. Because Coelophysis is one of the earliest 
theropods, its narrow pubis was thought to be diagnostic of 
Theropoda. However, this apomorphy is diagnostic of most, if 
not all, ceratosaurs, and a relatively broader pubis appears to 
be the ancestral condition for Theropoda (e.g., .■illosaurus. Mad- 
sen 1976). 

A more complete discussion of the evidence supporting 
monophyly of Ceratosauria will be presented elsewhere (Rowe, 
in prep.). 

Carnosauria Huene, 1920 (n. comb.) 

Temporal Range. — late Jurassic to late Cretaceous. 

Included J kx\.—. -Illosaurus fragdis. .icrocamhosaunis alokcnsis. Indosaurus 
matleyi. .4leclrosaiirus olseni. Drypiosaurus aqudungnis. .ilhenosaunis sarcoplia- 
giis. -i hlnatus. .-I. lancensis. .ilioranms remotiis. Dasplelosaurus torosus, Indo- 
suchiis raplorms. Tarhosaiirus hataar. and Tvrannnsaunts re.x. 

Diagnosis. — Based primarily upon the series of Allosaunis 
fragilis as described by Madsen (1976) and .-llheriosaunis libra- 
tits as described by Lambe (1917) and Russell (1970). These 
data were supplemented by personal examination of Allosaiints. 



10 



ORIGIN OF BIRDS AND EVOLUTION OF FLIGHT 



Albertosaurus, and Tyrannosaunis. and the descriptions of car- 
nosaurs published in Marsh (1896), Osborn( 1905, 1906, 1912, 
1917), Gilmore (1920), Matthew and Brown (1922), Janensch 
(1925), Sternberg (1932), Stovall and Langston (1950), Rozh- 
destvensky (1958, 1965), Walker (1964), Colbert and Russell 
( 1 969), Ostrom ( 1 969a), Steel ( 1 970), Gallon and Jensen (1979), 
and Barsbold (1983). 

The medium- to large-sized theropods such as Megalosaiirits* 
and Euslrcptospondyliis* possess some camosaurlike attributes. 
These taxa are examples of a pervasive problem in theropod 
phylogeny, namely, the "megalosaur" problem. Megalosaums* 
was the first dinosaur described, but it is represented by limited 
material with no diagnostic features distinguishing it from other 
large theropods. As the name implies, "megalosaurs" are larger 
theropods, and several of their apomorphies are probably size- 
related in that they are also seen in large omithischians and 
sauropodomorphs (e.g., femur longer than tibia). In view of 
profound character discordance, it is more parsimonious to ac- 
cept these shared apomorphies as examples of convergence be- 
tween "megalosaurs" and large omithischians or sauropodo- 
morphs. When considering the "megalosaurs" and Camosauria, 
however, the problem of distinguishing homology from con- 
vergence is more difficult. Camosauria shares many apomor- 
phies with a portion of Theropoda that includes extant birds, 
and these can hardly be considered size related (see below). The 
problem with "megalosaurs" is that they either do not have 
these apomorphies, or the appropriate portions of their skeletons 
are unknown. S. P. Welles is currently involved in a revision of 
the "megalosaurs," and until he has revised the alpha taxonomy 
of this confusing group of fossils, there is little point in consid- 
ering them further. 

Several camosaur apomorphies listed below are also present 
in other medium to large theropods such as Dilophosawus and 
Ceratosaunis. Among the size-related apomorphies are opistho- 
coelous cervicals, the greater length of the femur relative to the 
tibia, a robust skeleton, and enlarged neural spines and trans- 
verse processes in the trunk vertebrae. These attributes are seen 
in all large saurischians. Nevertheless, the taxa here included in 
Camosauria possess corroborating synapomorphies in addition 
to those related to their size, and other large theropods, such as 
Ceratosaunis and Dilophosaurus. do not. 

Camosauria is distinguished from other Theropoda consid- 
ered in this analysis in that it possesses the following synapo- 
morphies: orbit dorsoventrally elongate and roughly key- 
hole-shaped (Fig. IG, H); supraorbital crests in fully mature 
individuals (Fig. IG, H); frontals and parietals narrow and very 
short; reduction of mandibular fenestra (Fig. IG, H); further 
reduction of dentary overlap onto postdentary bones and man- 
dibular symphysis (indicating improved intramandibular joint, 
Romer 1956); pronounced development of bony shelf below 
mandibular condyle on lateral surface of surangular, presumably 
associated with insertion of enlarged pterygoideus musculature 
(Fig. IG, H); ilium expanded anterodorsally (Fig. 5D); strongly 
opisthocoelous cervical and anterior trunk vertebrae (conver- 
gent in penguins); digits II and III reduced in hand, especially 
the latter, which is shorter than digit I (Fig. 4L; analogous con- 
dition in oraithurine birds, but resulting from loss of phalanges); 
very robust postcranial skeleton with stout, relatively thick- 
walled long bones, shortened and stoutly constructed tmnk and 



cervical vertebrae (especially in tyrannosaurids), and large neu- 
ral spines and transverse processes throughout vertebral col- 
umn. 

Tyrannosauridae, including Albertosaurus. Tarhosaurus. and 
Tyrannosaurus. are further derived within this assemblage in 
that they have the following synapomorphies: lacrimal excludes 
frontal from orbit (Currie. in press a. b)\ enlarged surangular 
fenestra and pteo'goideus shelf (Fig. IH); ventral process of 
squamosal nearly honzontally oriented (Fig. IH); postorbital 
and jugal massive and anteriorly directed postorbital reentrant 
into orbit (Fig. IH); tooth row fails to reach posterior to antor- 
bital fenestra (Fig. IH); forelimb less than one-quarter of hind- 
limb length; wrist bones very reduced (convergent in omitho- 
mimids); third manal digit reduced to no more than metacarpal 
splint; ascending process very broad, extends dorsally for nearly 
one-third height of astragalus + tibia (convergent in coeluro- 
saurs); calcaneum very reduced; and proximal end of metatarsal 
III strongly constricted between metatarsals II and IV (conver- 
gent in omithomimids, elmisaurids*, and Hulsanpes). 

Ornithomimidae Marsh, 1890 

(n. comb.: includes Deinocheiridae of 

Osmolska and Roniewicz, 1970) 

Temporal Range. — late Jurassic to late Cretaceous. 

Included Taxa.— Elaphwsauriis bambergi. Archacornithommms asiaticus. 
Ornithomiinus edmonticus, O velo.x. O sedens. Strulhiomimus alius. Dromi- 
cciominms brevitcrtnis. D samiieli. Gallimimiis hullalus. Ingenta yanshim. Ga- 
ludimimia brcnpcs. and DcituniiciiKs ininficis. 

Diagnosis. — Based primarily upon Gallimunus bullalus as 
described by Osmolska et al. (1972) and data derived from 
Russell (1972). Additional evidence derived from personal ob- 
servation of Strulhiomimus and descriptions in Marsh (1890, 
1896), Osbom (1917), Parks (1928,1933), Gilmore ( 1 920, 1 933), 
Janensch (1925, 1929), Stemberg (1932, 1933, 1934). Ostrom 
(1969u, b. 1970, 1974a, 1976^)), and Barsbold (1983). The di- 
agnosis below is based on Upper Cretaceous omithomimids, 
although more complete knowledge of Lower Cretaceous and 
Upper Jurassic taxa may alter it. 

Comified beak as indicated by form of unworn margins of 
edentulous, beaklike jaws (Fig. II; convergent in Caenagnath- 
idae and modem birds); premaxilla enlarged and beaklike, 
broadly contacting nasal to exclude maxilla from extemal naris 
(Fig. 1 1); secondary palate formed by premaxillae and maxillae; 
reduced jugal and ventrally elongate postorbital; reduced lower 
temporal fenestra; quadrate strongly inclined so that distal end 
lies far forward of proximal end; bulbous parasphenoid (also in 
Troodontidae, and to a lesser extent in birds); metacarpal I 
elongate and all digits of subequal length (Fig. 4M); carpus very 
reduced with poorly defined articular facets on individual car- 
pals; terminal unguals less trenchant and recurved, with reduced 
basal tubera (suggesting loss of raptorial function for the hand); 
humerus lightly constructed and deltopectoral crest reduced; 
ischium ventrodistally recurved (Fig. 5E); metatarsus nartow, 
and elongate compared to tibia length; metatarsal III strongly 
pinched between metatarsal II and IV, barely contacting distal 
tarsals (analogous modifications of the metatarsus arose con- 
vergently in tyrannosaurid camosaurs, elmisaurids*, omithu- 
rine birds, and troodontids); pedal digits short and stout. 



GAUTHIER: SAURISCHIAN MONOPHYLY 



II 



Deinonychosauria Colbert and Russell, 1 969 

Temporal Range— early to late Cretaceous. 
Included Taxa.— Troodontidae and Dromaeosauridae. 

Diagnosis. — Modifications of the foot in general, and the 
second pedal digit in particular, indicate a raptorial function for 
the pes in Deinonychosauria (Colbert and Russell 1 969; Ostrom 
1969a. b). According to these authors, the subequal lengths of 
pedal digits III and IV, together with modification of the rap- 
torial pedal digit II, indicate functional didactyly during loco- 
motion. The ungual on pedal digit II bears a large, compressed, 
trenchant, strongly recurved, scimitarlike claw. The second pha- 
lanx is shortened and subequal to the first phalanx in length. 
Moreover, the second phalanx has a prominent heel postero- 
ventrally, and its anterior and posterior articular surfaces allow 
increased digital excursion. Troodontids may not be the sister- 
group of dromaeosaurs (see Section V). This point is not clear, 
however, and following previous authors, this taxon is accepted 
on the basis of the shared apomorphic resemblances in their 
feet. 

It is interesting to note that Osmolska (1982) argued that the 
form of the metatarso-phalangeal joint indicated that the second 
pedal digit functioned differently in troodontids (=sauromi- 
thoidids) and dromaeosaurs. This observation alone cannot be 
taken to indicate nonhomology, because the morphology of one 
could be a transformation of that seen in the other, or both 
could be transformations of some more general condition shared 
by their common ancestor. Troodontids vary in the degree to 
which the second pedal digit is modified (Russell, pers. comm.). 
For example, Troodon has a more specialized raptorial second 
pedal digit and is more like dromaeosaurs in this respect. How- 
ever, the second pedal digit is less modified in other troodontids 
(Osmolska 1982; Barsbold 1977). The possible effects of age, 
size, and sex on the degree of development of these characters 
has yet to be determined. Information on the possible influence 
of these factors might be gained from extant cariamids, Chunga 
and Cariama. that have analogously modified second pedal dig- 
its; examination of the biological roles of their feet may also 
provide some insight into the function of the second pedal digit 
in Deinonychosauria. (Although the information was received 
too late to include in this analysis, Currie has informed me that 
there is some evidence for a possible troodontid-omithomimid 
group; in light of this it would have been more appropriate to 
consider three separate basic taxa— 1 Dromaeosaurus. 2 Dei- 
nonychus-Velociraplor. and 3 Troodontidae— rather than one, 
viz., Deinonychosauria.) 



1982), Russell and Seguin (1982), Currie (in press a. b), and 
Wilson and Currie (in press). 

Barsbold ( 1 974) separated 5. junior from S. mongoliensis be- 
cause the former is 1.3 times larger, has a few more teeth, and 
the specimens derive from different stratigraphic formations. 
More specimens may indeed reveal that they are different taxa. 
However, the dilTerences between these specimens could reflect 
size and age; current evidence cannot exclude the possibility 
that S. junior is merely an adult of the smaller S. mongoliensis 
(Currie, in press a. b notes a similar size range for Troodon). 
Accordingly, these taxa will be considered synonymous in the 
following analysis. 

Anteromedially inclined orbits, suggesting broadly overlap- 
ping visual fields; deep depression in braincase in region of 
middle ear cavity (see Currie, in press a); bulbous parasphe- 
noidal rostrum (also present in omithomimids and in a less 
modified form in some birds); small, closely spaced teeth with 
enlarged (also seen in some dromaeosaurs), distally hooked den- 
ticles on posterior margin, anterior denticles reduced or absent 
at least in the lower jaw; deep, narrow Meckelian fossa of den- 
tary; additional caudal vertebrae incorporated into sacrum (six 
sacral vertebrae); rodlike metatarsus with proximally attenuate 
metatarsal III wedged between slender metatarsal II and robust 
metatarsal IV; metatarsals II and IV in contact anteriorly in 
front of proximal end of metatarsal III (also in elmisaurids* and 
omithomimids); and tonguelike distal articular surface of meta- 
tarsal III. 

Dromaeosauridae Matthew and Brown, 1922 

Temporal Range.— early to late Cretaceous. 

Included Ta\a. — Dminaeosaiinis alberiensis. Dciiwnychus annrrhopus. I'e- 
locirapror mongoliensis, and Adasaurus mongoliensis. 

Diagnosis. — Based primarily upon Deinonychus as described 
by Ostrom (1969a. b. 1974ft, 1976/'). Supplementary data de- 
rived from descriptions in Matthew and Brown (1922), Osbom 
(1924ft), Colbert and Russell (1969), Barsbold (1976, 1977, 1979, 
1983), Sues (1977, 1978), Bonaparte and Powell (1980), and 
casts of the plastotype of Deinonychus. Sues (1978) included 
Saurornitholestes* in Dromaeosauridae. but he did so on the 
basis of plesiomorphic resemblances; until the evidence sup- 
porting this placement is made explicit, this taxon will be con- 
sidered separately. 

Prezygapophyses and haemal arches exceed length of caudal 
vertebrae (Fig. 2G); short metatarsus compared to femur length; 
peculiar ginglymoid structure of the distal ends of metatarsals 
II and III; deeply grooved distal ginglymus of metatarsal II. 



Troodontidae Gilmore, 1924 
(=Saurornithoididae Barsbold, 1974) 

Temporal Range. — late Cretaceous. 

Included Ja\a. —Saurorniihoides mongoliensis. S. junior, and Troodon for- 
inosiis [=.Stenonychosaiiriis inequalis and Pecunodon bakkeri following Cume (in 
press h). 

Diagnosis. — Based upon personal observation of casts of 
Stenonychosaurus inequalis and descriptions and comparisons 
of both this taxon and Saurorniihoides in Osbom ( 1 924ft), Stem- 
berg (1932), Colbert and Russell (1969), Russell (1969), Bars- 
bold (1974, 1977, 1979, 1983), Sues (1978), Osmolska (1981, 



Avialae (n. txn.) 

(L.: avis, bird; alae. wings) 

Temporal Range. — late Jurassic to Recent. 

Included J axa.— .Archaeoplery.x lithographica* plus omithurine birds. 

Diagnosis. — Based primarily upon Archaeopteryx lithogra- 
phica* as described by Heilmann ( 1 926), de Beer ( 1 954), Ostrom 
(1972, 1973, 1974a. ft, 1975a. ft, 1976a), Wellnhofer (1974), 
Tarsitano and Hecht ( 1 980), Martin ( 1 983a, ft). Whetstone (1983) 
and upon personal observation of the Eichstatt specimen and 
casts of the London and Berlin specimens. 

This new taxon, Avialae, is named so as not to violate the 



12 



ORIGIN OF BIRDS AND EVOLUTION OF FLIGHT 



classificatory conventions of this work, in which widely used 
names hke Aves are restricted to living taxa in order to maxi- 
mize stability and phylogenetic informativeness. Because of 
feathers and the presumed ability to fly. Archaeopteryx* has 
always been considered a bird. This informal usage has been 
maintained above, and use of the informal term "bird" for this 
taxon will be continued in the following discussion. In a formal 
sense, however, "birds" and Aves will not be synonymous. The 
"wmged iheropods" included in Avialae possess the following 
synapomorphies distinguishing them from other Theropoda. 

Premaxillae elongate, narrow, and more pointed anteriorly, 
with longer nasal processes; maxillary process of premaxilla 
reduced so that maxilla participates broadly in external naris 
(also in troodontids; Currie, in press a); enlarged brain/basicra- 
nium (temporal musculature fails to extend origin onto frontal 
bones); double-condyled quadrate displaced from distal position 
on opisthotic to more anteromedial position in contact with 
prootic (Currie, pers. comm. and Walker, pers. comm., disagree 
with Whetstone's interpretation of the quadrate; Currie notes 
the anterior displacement of the quadrate in troodontids, and 
Walker does not consider the quadrate to be double-condyled 
in Arcliacnpiery.x*): maxillary and dentary teeth reduced in size 
and number (or lost), with unserrated crowns and enlarged roots 
that completely enclose replacement teeth within them (see 
Howgate 1984, for an alternative view); robust furcula for hy- 
pertrophied flight musculature (Olson and Feduccia 1979); scap- 
ula with more or less prominent acromion process for ligamen- 
tous connection to clavicle (see Martin 1983/). for alternative 
view); length/breadth ratio of scapula at midlength exceeds nine 
(not in penguins) and scapula tapers distally; acrocoracoid tuber- 
osity larger than in other coelurosaurs; coracoid enlarged and 
inflected posteromedially more so than in other coelurosaurs; 
very long forelimbs and hands (e.g., in Archaeopteryx* forelimb 
is 120-140% of hindlimb length, and more than twice as long 
as distance between glenoid and acetabulum), with forearm more 
than 87% of humerus length and metacarpal II approaching or 
exceeding one-half of humerus length; ischium compressed and 
dorsoventrally deep; compared to other theropods, tibia, fibula, 
and metatarsals relatively more elongate with respect to femur, 
regardless of body size (metatarsals short in penguins and some 
other birds, J. Cracraft, pers. comm.); fibula attenuate distally, 
and may not extend to end of tibia; proximal tarsals fused to 
tibia-fibula and to one another in adults; distal tarsals and meta- 
tarsals fused at least distally in fully adult individuals (conver- 
gent in some ceratosaurs, elmisaurids*. and Hulsanpes); first 
pedal digit elongate and reversed (may be reversed in some 
extant birds, R. Storer, pers. comm.), metatarsal 1 attached on 
distal quarter of metatarsal II; tail reduced to no more than 23 
free caudal vertebrae; feathers cover limbs and tail, feathers on 
lateral margins of tail and posterior margins of arms enlarged, 
curved, and asymmetrically vaned, indicating aerodynamic 
function (e.g., Feduccia and TordofT 1979). 

It is not certain that feathers are confined only to avialans 
among coelurosaurs. Compsognalhus apparently lacks them 
(Ostrom 1978), so feathers appear to have been absent in coe- 
lurosaurs ancestrally. However, Conipsognathiis is the only non- 
avialan theropod that is preserved in an environment of de- 
position conducive to the preservation of feather impressions. 
Thus, future finds may demonstrate that feathers arose prior to 
the origin of birds. 



As the use of the asterisk indicates, Archaeopteryx lithogra- 
phica* is here considered to possess no apomorphies that would 
not be expected in the common ancestor of all birds. Thus, the 
specimens referred to this taxon are placed here because of their 
geographic and stratigraphic occurrence and overall similarity. 
These specimens probably represent a single species, but such 
opinions should always be distinguished from those based on 
appropriate evidence. Notwithstanding the interesting possibil- 
ities suggested by Martin (1983/») and Howgate (1984), there is 
no unambiguous evidence indicating either paraphyly or mono- 
phyly, and these specimens will be referred to collectively as 
the metataxon Archaeopteryx lithographica*. Because of its gen- 
eralized morphology and stratigraphic position, the specimens 
of Archaeopteryx* could be parts of an ancestral population that 
gave rise to all later birds. Hypotheses of ancestral status can 
only be weakly supported in that they are based on negative 
evidence. Nevertheless, there is no unequivocal evidence sup- 
porting the alternative hypothesis, that Archaeopteryx* is 
monophyletic and thus not an ancestral bird. 

Whetstone (1983) and Martin (1983a) suggested that, com- 
pared to other birds, the squamosal is either reduced or absent 
in Archaeopteryx*. This interpretation is open to doubt in view 
of the preservation of the specimens with cranial material; each 
of the specimens was preserved such that upon separating the 
slabs, the skulls fractured between the main body of the skull 
and the lightly constructed elements surrounding the orbit and 
temporal fenestra. These authors contended that the squamosal 
was absent because there is no evidence for its sutural connection 
to the skull. This contention loses much of its force because 
these sutural surfaces are also absent in theropods in which the 
squamosal is known to be present (e.g., Syntarsits: M. A. Raath, 
pers. comm.). Under such circumstances, it is diflicult to dis- 
tinguish between absence and nonpreservation. If further finds 
corroborate the Whetstone-Martin hypothesis, then Archaeop- 
teryx* must be removed from metataxon status and its hy- 
pothesized ancestral position must be rejected. 

In keeping with one of the goals of this work, namely to 
provide a relevant series of outgroups for phylogenetic analyses 
among the major groups of extant birds, two more taxa within 
Avialae will be defined and diagnosed below. They are not basic 
taxa in this analysis, but it is necessary to consider Omithurae 
and Aves at this point because the concepts represented by these 
names as used in this study may differ from those employed by 
others. 

Omithurae is defined here in keeping with its original intent 
as a taxon encompassing all extant birds, as well as all other 
birds that are closer phylogenetically to extant birds than is 
Archaeopteryx*. Having been supplanted by Neomithes (Ga- 
dow 1 893), Omithurae (Haeckel 1 866) is seldom used in current 
ornithological literature; the obscurity of the name has saved it 
from the diversity of meanings that possible altemative names 
have developed, and Omithurae is thus an appropriate name 
for thi